ARAG » 8 9 ro Nr A dre met mr Bnr omkeer tn me nennen revert Bredere Prt et Ones ee tn Med en mn des de € a me de An IE NE ond me En arn nd ende sele TE. Lunar 0 zetten Ont = Em preda AP à en eee eee Se Bn NV De inn ond 2 Prenen wad lend By Sith boe vn Ent dnd nn ETA Oee D MT a n mtd nn nn dt Sint ate reek deri mt Orte pet die a nee" Menig nr dn II] er ner a win Rae wD odode Co Spas ee ree enne OAs TARA De OLE NOR Ee bp tee bree Ze Fn On bored BLEND deelne e Cera ot rn ee dee de tiv ih o a Li Eu Ca trenton Boye BY ‘ do des ereen Un 49 U ir ‘odo dn Bite ein bs I pee en eeb banden onp ii e tip Er DIET mand annen ne) sn mndée 2 Bun dotto br ELE imm ee dop cern e Bedi eenden OS md scene bee mahnte Oden Oth Er manen dif ot a te Ma Ph i Materia i inie ae rey ng are Melia ame ne en a in a © dtm Ent ad eut En tin È RAT AL ann tears: caste Pesce chemin del Hees Om Ba whe and Si sate ont On En esis rt nd en dat Edn ohn Phat i ene malin VE PT Pe TIA dees Uo Auer Pre ED re o> de ie TTI DL Fonda 4 PIERRE IE are COTE mom ets ON Fat CISTITE ane € terre erin nde votre Der pede ty pere De Or ren em ur n ae = pa nda tn rn let on nnen se ee LE nam ei De De oe Nair pae D mt DEAN EEn Rip Pest deren On Dench Ane Rita dre 7 Te nn DE Da a ee EC PELLE ETES a De A Eh tn en À md en em Antenne an amen nnn BA A do nn in cron nn Dein eee ee nun da ce Denn min nn nn Li I En rn nm nn er Te wdn in De Ae nenten dno np send Me E e Pai Ate Siw eM fim inthe SA Bre in eo too mente ope nan Ho dt Dd pn am nnn te rep ennen 20 En Bin ri Han Fe Te ie dn eN Medline, 6 29 8e Ben dn dn ine HO e PRONTI med bernard meth Fin D On te he ere fi EP Vale A en bin Zw ELON MLT AIA Bin di wende I E tin soe than meenden, wh an jn pin eri Bd Bate Met an ren rete fiv Sudato Brite Dm rer 8e une ans mente ne De ae mine en een denne Omer € Beene nennen ar Sue Di rm ermm Ata TALI ANA 8 R WETS AVA E ANT word SOTTO Ana dista EN beren enter On he bale © ard ents aten SER gn Aerien dn an en Eni pe Ata oen tn betonen Me eta nnen and OAN TETE ent sa event EIN tn ON D vee en VAM Sr mn ann VE Und PEEL UNI A CCE CNRS et AA A man IONICA Nenn mn ann "Nor eth ré een Ne reren ETES TETE D AREA ER A NEY VU EL EN, TA vin NAAN EEA U AR CLA Ae Où no DOUTE mind RITTER Fes Ut ave € enen Bhs Fa EN men nm Heres “a datent 2 È a Doper En an ALD er An EISEN Oe TU DIEU te N ni INNI DONNE EEE Beter an man none vern Iene AFIN er Ennn INTENTO WAN yay ee # DOTE RENTE EU farina {het DENTS vrind mee EVN rant att, ma arne nen end NA m m ARE mar kennen Lg Biere TETI en en ere ern een Era nn A am AN SX tn a NFTCORT DEAN valet, peet e Fyne Vere Ae Re rd ven en DORE SARA PUB A PA A eo NASA DOWNS As nti DORA Tale VX à a sf wien, DER TEEN TE DC Sm ETS Mara zione te: dm Rete mirata vos Ar Borne A on > x Sn RNA ah Ar Ce ARIA e ee SE 408 de A SAR CL N, UA Eur mon NOS ROTTA VINES Nodes een Peewee Pe Ra VR Ra PN Sr, A mm 919 ARR Fa 620 19 ia Romans My han Kurs Mn SITY MANY pirates tuin ‘i AT mn A A Ahr WIM OA an ME IAT De ans rt. DONEREN AAA iden. Par A À ein der eha D AN RY Art ORE NESTA A beer ter Co NE Dy mem Alea Rhy Me Mes, nme ay BeBe mini Fran ur Mn IR md Metre mu Sermon Me An Kirk ringe AT NR Race ante ee er VARA IN a AN eN Aen prenten gt RMT I NN urea MA te ENTO D on re NN Ben se OMA RG Ml ar mio Sar am aha PA nnn, nn emt Semele HARVARD UNIVERSITY È Library of the Museum of Comparative Zoology Volume 134, no. I, 1991 Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Published by the Nederlandse Entomologische Vereniging si Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 95i7, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zodlogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, ’s-Gravenhage C. GIELIS National Museum of Natural History, Leiden MCZ LIBRARY JUL 5 1 1991 PTEROPHORUS SCHAFFER IN THE ETHIOPIAN REGION (LEPIDOPTERA: PTEROPHORIDAE: HARVARD PTEROPHORINAE): NEW SPECIES AND CHECKLIST UNIVERSITY Gielis, C., 1991. Pterophorus Schiffer in the Ethiopian region (Lepidoptera: Pteropho- ridae: Pterophorinae): new species and checklist. - Tijdschrift voor Entomologie 134: 1- 8, figs. 1-21. [ISSN 0040-7496]. Published 1 July 1991. Pterophorus dallastai sp. n., P. uzungwe sp. n. and P. massai sp. n. are described. A provisional checklist of the Ethiopian Pterophorus species is given, with notes on all species. A lectotype is designated for P. cleronoma (Meyrick). P. centrocrates (Meyrick) is synonimised with P. rhyparias (Meyrick). The status of P. endophaea (Meyrick) and P. bacteriopa (Meyrick) is discussed. Correspondence: C. Gielis, Mr. Haafkensstraat 36, 4128 CJ Lexmond, The Netherlands. Keywords. - Lepidoptera; Pterophoridae; Prerophorus; new species; Africa. When the opportunity arose a number of Ethi- opian Pterophoridae from the collections of the Zoological Museum of the University of Copenha- gen (ZMUC), the Museum National d’Histoire na- turelle, Paris (MNHN) and the Royal Museum of Central Africa in Tervuuren (MRAC) were exam- ined. The results are here combined with data ob- tained from the collection of the British Museum of Natural History, London (BMNH), the Instituut voor Taxonomische Zoologie, Amsterdam (ZMAN) and the authors’ collection (CG). Some of the spe- cimens resembled, on superficial examination, pa- laearctic species. A closer examination of these species, however, showed a considerable variabil- ity, especially in the genitalia. This was particularly so in the genus Pterophorus, the subject of this publication. The species have usually poorly marked wings, which show a colour, ranging from purely white to pale grey-brown. Especially in the group of purely white specimens examination of the genitalia is essential. It is to be expected that more species will be recognized if more material from this region becomes available. The genus Pterophorus has its main distribution centre in the Palaearctic region. A high number of species (approximately 60) is known from West and Central Europe, the Mediterranean area, Tur- key and adjacent areas. From there the genus has a continuous distribution into the Southeast Asiatic region. Apart from Africa, further a few species are recorded from the Pacific area and the new world. Literature on the african representatives of the genus Pterophorus is scattered over a few smaller publications and mainly comprises single descrip- tions of species, without illustrations of externals or genitalia. Bigot (1969), in his review of the collection of MRAC, describes and illustrates the specimens in this collection, in this way fixing the new species described by him accurately. Since then, Bigot has given his attention mainly to the fauna of Madagascar. Only Arenberger (1986) and Gielis (1990) have published since on the Ethio- pian fauna, but on other subfamilies: Agdistinae and Platyptiliinae. Provisional checklist of Ethiopian Pterophorus Pterophorus Schäffer, 1766 Aciptilia Hiibner, [1825] Merrifieldia Tutt, 1905 Porrittia Tutt, 1905 Alucita auct., nec Linnaeus, 1758 1. albidus (Zeller, 1852) comb. n. 2. rhyparias (Meyrick, 1907) comb. n. centrocrates (Meyrick, 1933) syn. n. . ischnodactyla (Treitschhe, 1833) endophaea (Meyrick, 1930) comb. n. bacteriopa (Meyrick, 1922) comb. n. ceraunia (Bigot, 1969) comb. n. lampra (Bigot, 1969) comb. n. N DW mw TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 8. spissa (Bigot, 1969) comb. n. 9. lindneri (Amsel, 1963) comb. n. 10. candidalis (Walker, 1864) comb. n. 11. massat sp. n. 12. uzungwe sp. n. 13. dallastai sp. n. 14. cleronoma (Meyrick, 1920) comb. n. TAXONOMIC TREATMENT Pterophorus Schaffer, 1766 Type-species: Phalaena Alucita pentadactyla Linnaeus, 1758 Diagnosis. - Forewing cleft from % or less. Top of lobes acute, without a terminal margin. R1 ab- sent, R4 and RS fused, M3 and CU? forked or fused. Hindwing without a scale-tooth. Male genitalia asymmetrical, with asymmetrical costal processes. Female genitalia with a blister-like bursa copula- trix in which no, one or two signa, an often pro- nounced vesica seminalis, and often absent apo- physes anteriores. 1. Pterophorus albidus (Zeller, 1852) comb. n. (figs. 1, 10) Aciptila albida Zeller, 1852: 397. Diagnosis. - The species is characterized by the faint yellow colour. Male genitalia with a hooked costal process in the left valve and a large club-like process in the right valve. The female genitalia have the junction between the ductus bursae and the bursa copulatrix in the shape of a bulging pro- trusion with numerous sclerotized ridges. Remarks. - This species has originally been des- cribed from South Africa. This species can be easily recognized by its yellowish-white to greyish-white colour and the minute punctuation. After studying the male (fig. 1) and female (fig. 10) genitalia, the species turned out to have a wide distribution into the Indo-Australian region. Specimens have been Figs. 1-9. Pterophorus spp., male genitalia. — 1, P. albidus (Zeller) Rep. of S. Africa, slide cG 1694 (ZMAN); 2, P. rhyparias (Meyrick), British East Africa, slide CG 1870 (MNHN); 3, P. ischnodactyla (Treitschke, 1833). Rep. of S. Africa, slide CG 1695 (ZMAN); 4, P. ceraunia (Bigot), paratype, Zaire, slide Bigot (MRAC); 5, P. lampra (Bigot), paratype, Zaire, slide Bigot (MRAC); 6, P. spissa (Bigot), Central African Republic, slide cG 1878 (MNHN); 7, P. candidalis (Walker), Ethiopia, slide CG 1879 (MNHN); 8, P. dallastai sp. n., holotype; 9, P. wzungwe sp. n., holotype. GIELIS: Ethiopian Pterophorus examined from: Republic of South Africa, Zim- babwe, Kenya, Zaire, Tanzania, Gambia, Cöte d'Ivoire, Vietnam, Indonesia, New Guinea. 2. Pterophorus rbyparias (Meyrick, 1907) comb. n. (figs. 2, 11) Alucita rhyparias Meyrick, 1907: 489. Lectotype 9 (here designated): Rep. of S. Africa, Transvaal, Pretoria (Janse), genitalia slide BM 17993 (BMNH) [examined |. Alucita centrocrates Meyrick, 1933: 425. Holotype &: Belgian Congo (Zaire), N.E. Kivu, La Mutura, iii (Sey- del), genitalia slide Bigot (MRAC) [examined]. Syn. n. Diagnosis. - The species is characterized by the pale ochreous-white colour with some small black dots along the costa and dorsum of both forewing lobes. Male genitalia (fig. 2) with a long and short costal process in the left valve and a large club-like process in the right valve. The female genitalia (fig. 11) shows a distinct crescent-shaped distal margin of the 7th tergite and a many folded ap- pearance of the ductus bursae. Variation. - The colour of the species ranges from grey-white to pale ochreous-white. The number of dots on the forewing may be reduced. Distribution. - Rep. of S. Africa, Zaire, Kenya, Tanzania. Remarks. - Since the type-specimens of P. rhy- partas and centrocrates are respectively a female and a male, the synonymy, suggested by Aren- berger (personal communication) could only be proven by additional material. The evidence was found in a small series, present in ZMUC, where both sexes are present. This species is difficult to separate from the next, P. ischnodactyla Treitschke, and can only be distin- guished by the distinct genitalia. 3. Pterophorus ischnodactyla (Treitschke, 1833) (fig. 3) Alucita ischnodactyla Treitschke, 1833: 223. Diagnosis. - The species is characterized by the pale ochreous-white colour and the small black dots along the costa and dorsum of the forewing. Male genitalia (fig. 3) with a widely forked costal process in the left valve and a hooked single process in the right valve. The female genitalia show an exca- vated shape of the distal margin of the 7th tergite, the ductus bursae is bulged out, with hardly any transverse ridges. Remarks. - The Zoological Museum, Amster- dam (ZMAN) holds a specimen from the Republic of South Africa, Bloemfontein, 30.1.1951 (Ee), ge- nitalia CG 1695. This specimen represents the sole verified African record of the species known to me. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 10-15. Prerophorus spp., female genitalia. - 10, P. albidus (Zeller), Tanzania, slide cc 4018 (zMuc); 11, P. rhyparias (Meyrick), Zaire, slide Bigot (MRAC); 12, P. ceraunia (Bigot), Benin, slide cG 1881 (MNHN), 13, P. lampra (Bigot), Côte d'Ivoire, slide CG 2178 (CG); 14, P. spissa (Bigot), paratype, Zaire, slide Bigot (MRAC); 15, P. candidalis (Walker), Côte d'Ivoire, slide CG 2182 (CG). This means an extension of the distribution from the Mediterranean area towards the south of Africa. 4. Pterophorus endophaea (Meyrick, 1930) comb. n. Alucita endophaea Meyrick, 1930:576. Distribution. - Mozambique. Remarks. - See under P. bacteriopa 5. Pterophorus bacteriopa (Meyrick, 1922) comb. n. Alucita bacteriopa Meyrick, 1922: 41. Distribution. - Tanzania. Remarks. - P. endophaea and P. bacteriopa have not been examined, since I was unable to locate the type specimens. The description of both species indicates a relationship with the difficult to separ- ate species P. rhyparias and P. ischnodactyla. A synonymy based on the descriptions is suggestive, but not proven yet. 6. Pterophorus ceraunia (Bigot, 1969) comb. n. (figs. 4, 12) Aciptilia ceraunia Bigot, 1969: 204. Diagnosis. - The species belongs to the group of white species, with distinct black fringe markings. Differs in male genitalia (fig. 4) in having an asym- metrical pair of costal processes in the left valve and an equal long pair of processes in the right valve. In female genitalia (fig. 12) in having long ductal, longitudinally ridged, segments in both the junctions to the bursa copulatrix as to the vesica seminalis. Antrum longitudinal. Distribution. — Zaire, Benin. 7. Pterophorus lampra (Bigot, 1969) comb. n. (figs. 5, 13) Aciptilia lampra Bigot, 1969: 204. Diagnosis. - The forewings are purely white with some black scales near the base and inner margin of the first lobe. Differs in male genitalia (fig. 5) in having a single costal process in the left valve; slightly longer than the valve and a single long costal process in the right valve which is slightly shorter than the valve, in female genitalia (fig. 13) in having an arched 7th tergite, with the lateral positioned antrum, laterally progressing via GIELIS: Ethiopian Pterophorus the short ductus bursae into the blister-like bursa copulatrix. Distribution. - Zaire, Gabon, Côte d'Ivoire. 8. Pterophorus spissa (Bigot, 1969) comb. n. (figs. 6, 14) Aciptilia spissa Bigot, 1969: 202. Diagnosis. - Forewings pure white, with small black spots at the inner margin of both lobes. The fringes show a distinct black and white pattern. Differs in male genitalia (fig. 6) in having two costal processes in the left valve, of which the shorter is “th of the length of the longer. In the right valve two costal processes of equal length and a small transverse, short hook; in female genitalia (fig. 14) in having the ductus bursae slightly longer than the ductus to the vesica seminalis. Antrum large, almost square. Distribution. - Zaire, Central African Republic. 9. Pterophorus lindneri (Amsel, 1963) comb. n. Aciptilia lindneri Amsel, 1963: 6. Diagnosis. - The species is shining white, with- out any dark scales. Bursa copulatrix without a signum. Male genitalia. - Unknown. Female genitalia. - See diagnosis. Distribution. - Ethiopia. Remarks. - I have not examined the type-spec- imen, since the description by Amsel (1963) is adequate. It indicates the absence of a signum, which character differentiates the species from all its allies, which have one or two rosette-like signa in their bursa copulatrix. 10. Pterophorus candidalis (Walker, 1864) comb. n. (figs. 7, 15-17) Aciptilus candidalis Walker, 1864: 948. Redescription. - Wingspan 15-28 mm. Head white. Palpae short, white and greyish at top. An- tennae white, shortly ciliated. Thorax, mesothorax, tegulae and abdomen white. Legs white. Spurs of unequal length. Forewings cleft from 74, colour white with a faint yellowish grey gloss. Sparse dark scales at the base of the cleft and near the apex of the first lobe. Fringes white, at dorsum of both lobes with two faint greyish interruptions. Hind- wings white. Fringes white, at dorsum of second lobe a greyish interruption. Male genitalia (fig. 7). - Valvae asymmetrical. In TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 21 Figs. 16-21. Pterophorus spp., female genitalia. - 16, P. candidalis (Walker), Chad, slide CG 2186 (cG); 17, Idem, Tanzania, slide cc 4020 (zMuc); 18, P. dallastai sp. n., paratype, slide CG 1851 (MRAC); 19, P. massat sp. n., holotype, 20; P. uzungwe sp. n., paratype, slide cc 4040 (ZMUC); 21, P. cleronoma Meyrick, parailecitotype, slide Bigot (MNHN). right valve a pair of saccular spines; the ventral spine twice as long as the centrally placed spine. The left valve with two saccular spines; the ventral spine abruptly ending in a slender, acute top. The dorsal spine slender, length “4th of the length of the ventral spine. Tegumen simple. Uncus slender, of moderate length. Vinculum arched. Aedeagus straight. Female genitalia (figs. 15-17). — Antrum in shape of a slender rectangular plate, with a rounded top around a small hole. Ductus bursae and ductus seminalis well-developed with longitudinal ridges. Bursa copulatrix vesicular, with a single rosette- like signum. Variation. - The female genitalia show variation in the length of the ductus seminalis. In specimens from Céte d'Ivoire (fig. 15) it is longer than the ductus bursae; from Chad and Ethiopia (fig. 16) of equal length, and in Tanzania (fig. 17) shorter than the ductus bursae. Distribution. — Ethiopia, Kenya, Tanzania, Zaire, Chad, Côte d'Ivoire, Sierra Leone. Remarks. — The type specimen from Sierra Leone (BMNH) is missing its abdomen. After com- paring with the specimens from areas relatively close to the type locality, I agree with Bigot (1969) about the identity of the species. 11. Pterophorus dallastai sp. n. (figs. 8, 18) Type material. - Holotype 4: Zaire [Congo Belge], P. N. A., Secteur Tschiaberimu, Mont Kamatondi pres Ki- zungu, 2850 m., 22.iv.1955 (Vanschuytbroeck & Fontein, no. 12785), genitalia slide cc 1850 (MRAC). — Paratypes: 1 9, Zaire [Congo Belge], P. N. A., Secteur Tschiaberimu, Mont Musimba près Musabaki, 2450 m., 25.11.1954 (Van- schuytbroeck & Synave, no. 7830), genitalia slide cc 1849 (MRAC); 2 9, Zaire [Congo Belge], P. N. A, Secteur Tschiaberimu, Riv. Musavaki, affl. Talya Nord, 2720 m, 26-29.111.1954 (Vanschuytbroeck & Synave, no. 7828-29), genitalia slides CG 1851, 1852 (MRAC, CG). Diagnosis. - The species is closely related to P. candidalis (Walker). It is characterized by the slightly ochreous tinge of the forewing, the saccular spines in the male genitalia and the shape of the antrum in the female genitalia. Description. - Male, female. Wingspan 24-28 mm. External characters as in P. candidalis (Walker). The colour of the forewings is slightly ochreous tinged. Male genitalia (fig. 8). — Valvae asymmetrical. Right valve narrow, lanceolate, with a long curved and a short stout saccular spine. Left valve widen- ing towards %th of valve length; with a long curved and a short, wavy saccular spine. Tegumen bilobate. Uncus slender, moderate. Vinculum narrow, GIELIS: Ethiopian Pterophorus arched. Aedeagus tube-like, towards top gradually narrowing. Female genitalia (fig. 18). — Antrum large, rounded, without a central hole. Ductus bursae longer than ductus seminalis, both ducti with lon- gitudinal ridges. Bursa copulatrix vesicular, with a single rosette-like signum. Variation. - The specimens show some variation in the intensity of the black scaling on the fore- wings. Biology. - The moths were collected in March and April. Immature stages unknown. Distribution. - Zaire. Etymology. - A noun in genitive case, named in honour of Dr. U. Dall’asta, curator of Lepidoptera in MRAC. 12. Pterophorus massai sp. n. (fig. 19) Type material. - Holotype 9: Kenya (Afr. Or. Angl.), Nairobi, Plaine Massai, viii.1904 (Alluaud), genitalia slide cG 1875 (MNHN). Diagnosis. - The species is closely related to P. candidalis (Walker) and P. dallastai Gielis, but char- acterized by the female genitalia: the ratio of the length of the ductus bursae and the ductus semina- lis is three to one. Description. - Female. Wingspan 25 mm. Exter- nal characters as in P. candidalis (Walker). Male genitalia. - Unknown. Female genitalia (fig. 19). - Antrum large, end- ing roof-like, with an oval hole. Ductus bursae three times longer than ductus seminalis, both with longitudinal ridges. At the junction of the ductus bursae and ductus seminalis two rows of small spines. Bursa copulatrix vesicular, with a rosette- like signum. Biology. - The moth was collected in August. Immature stages unknown. Distribution. - Kenya. Etymology. - Massai, a noun in apposition, named after the type locality. 13. Pterophorus uzungwe sp. n. (figs. 9, 20) Type material. - Holotype &: Tanzania, Uzungwe Mts., Mwanihane Forest above Sanje, 1000 m, 1.viii.1981 (Stoltze & Schaff), genitalia slide cc 4039 (ZMUC). Par- atype 9: same locality and data, genitalia slide cG 4040 (GS, Diagnosis. - The species is characterized by the shape of the saccular arms in the male and the double signum in the female. The size is relatively TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 small for species of its group. Description. - Male, female. Wingspan 18-19 mm. External characters as in P. candidalis (Walker). Male genitalia (fig. 9). - Valvae asymmetrical. Right valve with a long, almost straight, saccular spine. Left valve with a curved saccular spine, with two small lateral thorns. Tegumen bilobate. Uncus slender, moderate. Vinculum rather wide. Aedea- gus tube-like. Female genitalia (fig. 20). - Antrum three times longer than wide, club-like, with a large central distal hole. Ductus bursae and ductus seminalis of equal length, both with longitudinal ridges. Bursa copulatrix with two rosette-like signa. Biology. - The moths were collected in August. Immature stages unknown. Distribution. - Tanzania. Etymology. - Uzangwe, a noun in apposition, named after the type locality. 14. Pterophorus cleronoma (Meyrick, 1920) (fig. 21) Alucita cleronoma Meyrick, 1920: 41. Lectotype 9 (here designated): Brit. E. Africa, Kenya, Mt. Kenya, i- ii.1912 (Alluaud & Jeannel), genitalia slide BM 17987 (BMNH) [examined]. Other material examined. — Paralectotype Q: locality and data as lectotype, genitalia slide Bigot (MNHN) [examined]. Diagnosis. - The species is characterized by the shape of the female genitalia. Especially the grad- ually narrowing antrum, in combination with the absence of a signum is diagnostic. Female genitalia (fig. 21). - Antrum gradually narrowing. Ductus bursae slender. Bursa copulatrix vesicular. No signum. Apophyses anteriores ab- sent. Apophyses posteriores three times longer than papillae anales. Distribution. - Kenya. Remarks. - In his description Meyrick did not designate a holotype. Both BMNH and MNHN ap- peared to hold a specimen labelled ‘type’, the BMNH specimen is herewith designated as lectotype. ACKNOWLEDGMENTS I wish to thank Mr. B. Brugge (ZMAN), Dr. U. Dall’asta (MRAC), Mr. O. Karsholt (ZMUC), Dr. G. C. Luquet (MNHN) and Mr. M. Shaffer (BMNH) for the opportunity to study the Pterophoridae in their museum, Mr. H. W. van der Wolf for correcting the English text and Mrs. van de Sant for typing the manuscript. REFERENCES Amsel, H. G., 1963. Kleinschmetterlinge aus Aethiopien. - Stuttgarter Beiträge zur Naturkunde 121: 1-12. Arenberger, E., 1986. Die Agdistis-Arten der Aethiopi- schen Region, 1. Beitrag. - Mitteilungen der Schwe- izerischen entomologischen Gesellschaft 59: 187-196. Bigot, L., 1969. Les Lepidopteres Pterophoridae du musee royal de l’Afrique Centrale, à Tervuren. - Revue de Zoologie et de Botanique Africaines 79: 165-206. Gielis, C., 1990. Neue Pterophoridae der Aethiopischen Fauna. - Entomologische Zeitschrift, Frankfurt 100: 113-125. Meyrick, E., 1907. Notes and descriptions on Pteropho- ridae and Orneonidae. - Transactions of the Entomo- logical Society of London 1907: 471-511. Meyrick, E., 1920. Microlepidoptera. - In: Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911- 1912). Résultats scientifiques. Lepidoptera, II: 33-120. Paris. Meyrick, E., 1922. New microlepidoptera of the German entomological institute. - Entomologische Mitteilun- gen 11: 44. Meyrick, E., 1930 (reprint 1969). Exotic microlepidop- tera 3 (18): 545-576. - Hampton, England. Meyrick, E., 1933 (reprint 1969). Exotic microlepidop- tera 4 (14): 417-448. - Hampton, England. Treitschke, F., 1833. Die Schmetterlinge von Europa 9: 1- 294. - Leipzig. Walker, F., 1864. List of the specimens of lepidopterous insects in the British Museum 30. Lepidoptera Heter- ocera: 926-953. - London. Zeller, P. C., 1852. Revision der Pterophoriden. - Linnaea Entomologia 1: 319-413. Received: 5 June 1990 Revised version accepted: 2 January 1991 R. V. HENSEN Utrecht, The Netherlands REVIEW OF MALESIAN SPHECINA (HYMENOPTERA, SPHECIDAE, SPHECINAE) Hensen, R. V., 1991. Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae). - Tijdschrift voor Entomologie 134: 9-30, figs. 1-65. [ISSN 0040-7496]. Published 1 July 1991. The species of Sphex and Isodontia (subtribe Sphecina) in the Malesian region are reviewed. Twenty species are recognized in Isodontia, which are segregated into five species-groups, while 21 species are recognized in Sphex, divided among three species- groups. Keys are provided to the genera, species-groups and species. Synonymy and distribution of the species is discussed. Twelve new species are described, viz. I. pilipes (Burma to Sumatra, Borneo), I. cestra, I. elsei (both Sulawesi), I. nidulans (Philippines), I immaculata, 1. capillata, I. delicata, I. papua, S. wilsoni, S. rex (all New Guinea), S. solomon (Solomon Islands), and S. walshae (NE India to Sumatra, Borneo). The following new synonymies are established: Sphex morosus Smith, 1861, S. volatilis Smith, 1861, S. triodon Kohl, 1890, S. abditus Kohl, 1895 and S. abditus nugenti Turner, 1910 are synonymized with Isodontia aurifrons (Smith, 1859); I. diodon alemon Van der Vecht, 1957 with I. severini (Kohl, 1898); S. wstulatus Kohl, 1890, S. simplex Kohl, 1898 and S. insularis Cameron, 1901 with I. praslinia (Guérin-Méneville, 1831); S. albohirtus (Turner, 1908) with I. sepicola (Smith, 1859) and S. fulvohirtus Bingham, 1890 with S. diabolicus Smith, 1858. Raymond V. Hensen, W.A. Vultostraat 147, 3523 TW Utrecht, The Netherlands. Key words. - Malesia, Oriental region, Australian region, Sphecidae, Sphex, Isodontia, taxonomy. In this paper the species of the subtribe Sphecina comprising the genera Sphex Linnaeus and Isodon- tia Patton are briefly treated as far as they occur in the Malesian area. The Malesian area is used here in a broad sense, i. e. comprising Southern Thai- land, Malaysia, Indonesia, the Philippines, New Guinea and the Solomon Islands. Sphex and Isodontia are large, active insects, and some of the species treated here are among the most commonly encountered digger wasps in the area. The last comprehensive publication dealing with the Malesian species was that by Kohl (1890), but the taxonomy of the group was updated in many smaller papers (eg. Kohl 1895, 1898, Van der Vecht 1957, 1973). Therefore, it is surprising that so many new species remained undetected. Furth- ermore, a considerable number of new synonyms were discovered in the course of this study. Comprehensive diagnoses of the morphology and biology of the two genera were given by Bohart & Menke (1976), and need not be repeated here. One of the aims of this study is to define and diagnose a number of species-groups. Most of these groups are considered monophyletic, being based on inferred synapomorphies. Others, in particular the Sphex subtruncatus-group are possibly para- phyletic. It seems premature to pay more attention to the phylogeny of the subtribe here, as a thorough treatment of this subject will require study of all species in these genera. This paper deals strictly with species: it does not present a classification below the species-level. Al- though past workers (eg. Van der Vecht & Krom- bein 1955) have recognized subspecies in the Male- sian Isodontia, especially for colour differences from island to island, I have not recognized them in this paper. Subspecific names recognized in the past are simply treated here as synonyms of the nominate form. On the other hand, several taxa regarded as subspecies in the past are elevated to species rank. This study was initiated by dr. J. van der Vecht, and in many instances I have relied on his notes concerning literature, type material and specimens. As far as type specimens are concerned, I have indicated examination by Van der Vecht with a single exclamation mark, by myself with a double exclamation mark after the type depository (e.g. BMNH!!). A question mark behind an abbreviation of a museum means that a type was looked for but not found in that particular museum. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 A few thousand specimens were examined in the course of this study. I have refrained from listing these explicitly, except for the type material of new species. The outlines of distribution, as given for each species are entirely based on examination of specimens by me or by Van der Vecht, except as noted. Abbreviations. - Names of museums in which types are preserved are abbreviated as follows. BMNH Natural History Museum, London, Eng- land. BRI Biosystematics Research Institute, Ot- tawa, Canada. BPBM Bernice P. Bishop Museum, Honolulu, Hawaii, USA. CAS California Academy of Sciences, San Francisco, USA. DEI Deutsche Entomologische Institut, Eberswalde, Germany. IRSN Institut Royal de Sciences Naturelles, Brussels, Belgium. MCG Museo Civico di Storia Naturale, Genova, Italy. MCZ Museum of Comparative Zoology, Cam- bridge, Massachusetts, USA. MNHN Muséum National d’Histoire Naturelle, Paris, France. NMB Naturhistorisches Museum, Basel, Swit- zerland. NMW Naturhistorisches Museum, Vienna, Austria. OUM Oxford University Museum, Oxford, England. RMNH Nationaal Natuurhistorisch Museum, Leiden, The Netherlands. TMB Termeszettudomanyi Muzeum, Budap- est, Hungary. USNM National Museum of Natural History, Smithsonian Institution, Washington, D. CAUSA UZMC Universitetets Zoologisk Museum, Co- penhagen, Denmark. ZMB Zoologisches Museum der Humboldt- universität, Berlin, Germany. The following morphological terms are abbre- viated. IOC Shortest interocular distance across the clypeus. IOV Shortest interocular distance across the vertex. ACKNOWLEDGEMENTS Dr. J. van der Vecht donated his personal notes and provided many of the specimens that formed 10 the foundation of this study. The following persons aided in various ways, including arranging loans of material, assistance and hospitality at my visits to various museums, etc. In alphabetical order: Dr. ing. C. van Achterberg, Leyden; Dr. J. M. Carpen- ter, Cambridge, Mass.; Dr. J. Casevitz-Weulersse, Paris; Dr. P. Dessart, Brussels; Dr. M. Fischer, Vienna; Dr. F. Koch, Berlin; Dr. ©. Lomholdt, Copenhagen; Dr. A. S. Menke, Washington DC; Mr. G. M. Nishida, Honolulu; Dr. J. Oehlcke, Eberswalde; Dr. C. O’Toole, Oxford; Dr. J. Papp, Budapest; Dr. W. J. Pulawski, San Francisco; Dr. M. Sharkey, Ottawa; Mr. C. R. Vardy, London. I wish to express my sincere gratitude to these persons, and others who took a constructive inter- est in my activities. Special thanks are due to Dr. A. S. menke for reviewing the manuscript. A visit to the Natural History Museum, London, and a collecting trip to Indonesia were supported by the Uyttenboogaart-Eliasen Stichting. SYSTEMATIC PART Key to genera and species-groups 1. Sides of propodeum with complete spiracular groove; basal vein of second submarginal cell as long as or longer than anterior vein; female with well developed tarsal rake, composed of long, flattened spines (fig. 62) (Sphex) ..... 2 - Sides of propodeum without spiracular groove; basal vein of second submarginal cell usually shorter than anterior vein; female without tar- salfrakenlsodontsa) tae CRE EN 4 2. Metanotum medially with pair of tubercles or with single tubercle, that is more or less im- Pressed/imediallyar ae RR SERBIEN ORA eh eo a group of S. argentatus (p. 20) - Metanotum without tubercles, flat or slightly CONVEXIMECIA VARE ee eae tea eee ee ee 3 3. Scutellum of female flat, shiny (sometimes densely covered with tomentum); male flagel- lum without placoids; claw teeth small, per- pendicularstorclawA(ti 0859) EUR ER ee group of S. resplendens (p. 22) — Scutellum of female convex, often with median furrow, usually dull; at least fifth flagellomere of male with placoid (figs. 43-45); claw teeth large, at sharp angle to claw (fig. 60)........... VELEN SEE group of S. subtruncatus (p. 26) AE AMES ATI 9197 IA NIIR WI DEE AO NOA 5 SHRemales Ent MNT LAME DIE ONE dana 9 5. Mandibles bidentate ............................ 6 — Mandibles tridentate ........................... 7 6. Mandibles very slender, nearly straight (fig. 8); inner orbits strongly converging below; petiole 10. 11. 19% longer than hind basitarsus; small, slender spe- cies, body length 13-19 mm ES eg A group of I. diodon (p. 14) Mandibles stout and arcuate (fig. 6); inner or- bits parallel or slightly diverging below; peti- ole shorter than hind basitarsus; large, stout species, body length 20-28 mm group of I. chrysorrhoea (p. 13) . Fourth flagellomere with well defined placoid (figs. 35-38) group of I. praslinia (p. o Flagellum without placoids . Basal inner tooth of mandible strongly jee loped, much stronger than second inner tooth (fig. 10); basal half of clypeus with blunt lon- Ritudinalicasina e een: group of I. ochroptera (p. 19) Basal inner tooth of mandibles smaller (fig. 2); basal half of clypeus at most with trace of longitudinal carina group of I. aurifrons (p. 11) . Widest part of frons slightly narrower than clypeus in the middle, inner orbits slightly di- verging below; outer side of hind tibia with more than 20 short spines; body length 29-35 mm group of I. chrysorrhoea (p. 13) Inner orbits parallel or converging below; outer side of hind tibia with at most 10-12 short spines; body length 14-26 mm Mandibles very slender, straight, apically bifid (fig. 7); hind tibiae with 1-3 short spines on outer side; fourth segment of hind tarsi wider than long eee to group of I. diodon (p. 14) Mandibles robust, curved, apically tridentate (fig. 9); hind tibiae with 5-10 spines on outer side; fourth segment of hind tarsi elongate ao agri ident Retest rae Beiden! 11 Labrum with single median longitudinal car- ina, that does not project apically; apical mar- gin of clypeus with pair of well differentiated teeth (fig. 9); pilosity of head and mesosoma black group of I. ochroptera (p. 19) Labrum apically with two parallel, raised, pro- jecting carinae; apical clypeal margin medially notched (fig. 11), or with pair of short teeth (fig. 1); pilosity variable First flagellomere distinctly longer than se- cond; antennae not or hardly longitudinally ridged (fig. 21); mandible stout (fig. 11) group of I. praslinia (p. 19) First flagellomere hardly longer than second (fig. 19), or (I. pilipes) mandibles slender (fig. 3); antennae longitudinally ridged (fig. 19) group of I. aurifrons (p. 11) HENSEN: Malesian Sphecina Genus Isodontia Paton Group of Isodontia aurifrons Diagnosis Body length 4 25-23 mm, 9 17-26 mm. Labrum of female with a pair of short parallel projecting carinae apically; mandible tridentate in both sexes, sometimes with weak third inner tooth near apex; clypeus of female with median notch; first flagel- lomere as long as second; flagellum of female us- ually with longitudinal sulci on second to tenth segment; third to seventh flagellomere in male with convex, papillate sensory areas; outer side of hind tibia with 5-9 short spines; petiole usually shorter than hind basitarsus. Species included Species treated. - I. aurifrons (Sm.), formosicola (Strand), pilipes sp. n. Other included species. - I. mazdli (Yasumatsu) from Japan. Key to species ile 2: Females dente ee Ce 2. MAESTRI I TIE TR 4 First flagellomere much longer than second (fig. 20); Burma, Sumatra, Borneo pilipes sp. n. First flagellomere not or scarcely longer than second (fig. 19) . Metasoma largely or entirely dull red; clypeus with median carina over most of its length; Taiwan, China, S. Thailand, Malaysia, Sumatra, Borneo formosicola (Strand) Metasoma entirely black; clypeus with short carina near base or medially; Sikkim to China and Malaysia, Malesian region, Queensland aurifrons (Smith) . Metasoma largely dull red; third flagellomere as long as second (fig. 23); clypeus apically with pair of short teeth; wings yellowish formosicola (Strand) Metasoma entirely black; third flagellomere longer than second; clypeus apically without teeth (fig. 2); wings brown or clear nN . Petiole as long as hind basitarsus; first flagel- lomere longer than second (fig. 24) pilipes sp. n. Petiole shorter than hind basitarsus; first fla- gellomere as long as second (fig. 22) aurifrons (Smith) 11 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 1-12. Isodontia, clypeus and mandibles. - 1, aurifrons, 9; 2, aurifrons, 6; 3, pilipes, 9; 4, pilipes, &; 5, chrysorrhoea, 9; 6, chrysorrhoea, 6; 7, cestra, Q; 8, delicata, 8; 9, ochroptera, 9; 10, ochroptera, Ô; 11, praslinia, 9; 12, praslinia, €. Isodontia aurifrons (Smith) (figs. 1-2, 19, 22) Sphex aurifrons Smith, 1859: 157. Lectotype 9 (present designation): Indonesia, Aru (OUM!!) Sphex morosus Smith, 1860: 122. Holotype @: Indonesia, Bacan (OUM!!) Syn. n. Sphex volatilis Smith, 1860: 122. Holotype 9: Indonesia, Bacan (OUM!!) Syn. n. Sphex triodon Kohl, 1890: 377. Lectotype @ (present designation): Indonesia, Java (NMW!!) Syn. n. Sphex abditus Kohl, 1895: 51. Holotype 9: Sikkim (NMW!!) Syn. n. Sphex abditus nugenti Turner, 1910a: 343, 345. Holotype Q: Australia, Cairns (BMNH!!) Syn. n. Distribution. - Continental Asia from Southern India and Sri Lanka to China and Malaysia, Indone- sia, Philippines, New Guinea, Queensland. Isodontia formosicola (Strand) (fig. 23) Sphex (Isodontia) formosicola Strand, 1913: 82. Lecto- type Q (present designation): Taiwan: Hoozan (DEI!!) Distribution. - Continental Asia from China to Malaysia, Taiwan, Sumatra, Borneo. Isodontia pilipes sp. n. (figs. 3-4, 20, 24, 46) Type material. - Holotype 8, Indonesia, 'S. Sumatra, + 600 m, Res. Benkoelen, Tandjong Sakti, 16-19 VII, M. E. Walsh leg. 1935’ (RMNH). - Paratypes. Burma: Tenasse- rim, Ihaungyin Valley, v.1893, ‘Sphex aurifrons Sm., @ (1 2), Tenasserim, Ataran Valley, xii.1891 (1 9), Burma, Middle Tenasserim, Haundraw Valley, viii.1894 (1 9), all C. T. Bingham (BMNH). Indonesia: S. Sumatra, Benkulen, 50 m, 19-21.v.1935, M. E. Walsh (1 6, RMNH); W. Suma- tra, Padangpanjang, 800 m, 1.v.1988, R. Hensen (1 8, RMNH). Malaysia: Sabah, Ranau 8 mi. N., Paring Hot Springs, 500 m, 8-11.x.1958, L. W. Quate (1 8, BMNH); Borneo, , v.1900, P. Cameron coll., ‘Sphex nigripes, Sm.’ (1 9, BMNH); Borneo, Kuching, v.1900 (1 4, BMNH). Diagnosis Within the aurifrons-group, pilipes is character- ized by the first flagellomere being distinctly longer than the second. Additionally, in the male, the petiole is as long as the hind basitarsus (shorter in the other species). I. pzlipes may actually be closest to the Japanese I. maidli (Yasumatsu). It differs from that species in the male by the presence of hair-fringes on the sternites, and in the relative lengths of the proximal flagellomere (1 : 1 : 1.4 in matdli). In the female I. pilipes is characterized by the dark hind tibiae (reddish in maid/1) and the presence of transverse rugae medially on the prop- odeum. HENSEN: Malesian Sphecina Description Female. - Body length 17.5-21.0 mm; length of forewing 12.5-15.0 mm. Black, mandible medially reddish, tegulae with margin brownish yellow or reddish, distinctly lighter than the center. Wings yellowish, apical margins and particularly apex of forewing infuscated. Erect pilosity of head and mesosoma brown but yellowish on genae, venter of mesosoma and propodeum. Face with rather sparse silvery tomentum, mesosoma without con- spicuous tomentum. Clypeus 0.9 times as long as IOC, with median carina on dorsal half; 10V 1.0 times IOC; relative lengths of flagellomeres I-III 1 : 0.8 : 0.85; mandibles slender. Mesoscutum shiny, sparsely finely punctate, punctures on disc 1-3 di- ameters apart; propodeal dorsum shiny, rugose- punctate, with short transverse rugae medially. Pe- tiole 0.8 times as long as hind basitarsus. Male. - Body length 15.0-17.0 mm; length of forewing 11.0-12.5 mm. Like the female, except for the following. Black; mesoscutum with faint greyish shine. Wings light brown, with violaceous shine. Erect pilosity of head brownish to black, of mesosoma yellowish grey, but darker on mesoscu- tum. Clypeus 0.85 times as long as IOC; IOV 1.0-1.1 times IOC (1.0 in type); relative lengths of flagel- lomeres I-III 1 : 1.0 : 1.3; flagellomeres III-VIII with convex, papillate sensory areas; mandible with faint third inner tooth near apex; propodeal dorsum finely punctate-reticulate. Petiole 1.0-1.05 times as long as hind basitarsus; fourth to seventh sternite with fringes of appressed brown setosity; gonostyle simple. Etymology Pilipes is a noun in apposition, from Latin pilus = hair and pes = leg. Group of Isodontia chrysorrhoea (Kohl) Diagnosis Body length 9 29-34 mm, @ 20-28 mm; imme- diately recognized by their large size, and the pres- ence of yellow bands on the metasoma. Female: labrum not modified; mandible tridentate; clypeus rounded; first flagellomere distinctly longer than second. Male: mandible bidentate; clypeus shal- lowly emarginate; fourth to sixth flagellomere with broad, convex papillate sensory area. Species included Species treated. - I. chrysorrhoea (Kohl). Other included species. - I. auripygata Strand from Taiwan, China and Vietnam. Isodontia chrysorrhoea (Kohl) (figs. 5-6, 25) 13 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Sphex apicalis Smith, 1856; 253. Holotype 9: Indonesia, Sumatra (BMNH!!) [non Smith, 1856: 262]. Sphex (Isodontia) chrysorrhoeus Kohl, 1890: 371 [new name for S. apicalis Smith, 1856: 253]. Sphex (Isodontia) hewitti Cameron, 1906: 119. Holotype 8: Malaysia, Sarawak, Kuching (BMNH!!). Distribution. - S. Thailand, Indonesia (Sumatra, Java, Borneo, Nias), Malaysia (Sarawak); Philip- pines (Mindanao). Group of Isodontia diodon (Kohl) Diagnosis Body length & 12-18 mm, 9 14-19 mm. Labrum of female simple; mandible abnormally slender, nearly straight, bidentate in male, the apex bifid in female and with small outer tooth; clypeus of fe- male simple; relative lengths of flagellomeres var- iable; flagellum of female without sensory areas; flagellum of male with flat placoids on one or more flagellomeres or with broad, convex, papillate sen- sory areas. Scutellum and metanotum convex; propodeal dorsum reticulate to transversely stri- gose; outer side of hind tibia with 1-3 spines. Pe- tiole usually longer than hind basitarsus. Species included Species treated. - I. capillata sp. n., cestra sp. n., delicata sp. n., diodon (Kohl), egens (Kohl), franz: (Kohl), maculata sp. n., papua sp. n., permutans (Turner), severini (Kohl). Other included species. - I. nıgelloides (Strand) from Sri Lanka and Southern India and an un- named Australian species. Key to species 1. Females (unknown: capillata, delicata, papua) =) Males (unknown: man) ee 8 2. Forewing with sharply differentiated dark markrapicallly SS. ri BERE D denti 3) - Forewing darkened apically, but dark area not Shacplyidelimitedkt eas. ned ea E 6 SE acialktomentumksolden: obi 4 =MFacialitomentumisilveryietfa st. dere 2: 5 4. Petiole as long as hind basitarsus; first meta- somal tergite black, laterally reddish translu- cent SUIAWESITe ra en. ker cestra sp. n. - Petiole longer than hind basitarsus; first met- asomal tergite red; Sumatra, Java, Borneo, Phi- lippines, Nusa Tenggara, Moluccas ............ severini (Kohl) 5. Metasoma, except petiole, entirely red; Borneo SERA CO Led RE franzi (Cameron) — Metasoma black, with at most first tergite and posterior margins of other tergites reddish; 14 10. 1% 122 15: 14. 157 diodon (Kohl) . Facial tomentum golden (except specimens from Gizo Isl.); New Guinea, Solomon Islands DIS DT NE permutans (Turner) Facialsıtomentum silyveryas ee dl . Mesosoma with conspicuous silvery tomentum on pronotal lobes, metanotum and sides of propodeum; New Guinea, Bismarck Islands, BAR ER ee n egens (Kohl) Mesosoma without silvery tomentum: Bis- marckilslands Her immaculata sp. n. . Flagellomeres I-II short, III at least twice as longsasıl(es- fis220) a ee ee 9 Flagellomeres I-II longer, III at most 1.5 times asilong:asil6 (ep e 2 14 . Flagellomeres III-VI with broad, flat placoids over their entire length (figs. 26, 30,31) 10 Flagellomere III at most with small apical pla- CONIC GS este PS2) TI 13 Posterior tergites with conspicuous, dense yel- low pilosity; gonostyles apicodorsally with sharply pointed process (fig. 47); New Guinea capillata sp. n. Facial tomentum silvery; posterior tergites with sparse white or yellowish pilosity; gonos- tyles apicodorsally rounded (figs. 52, 54) or Withibroadiangle (fig OE 11 Mesosoma without tomentum; gonostyles apicodorsally swollen and rounded (fig. 52); Bismarck Islands... immaculata sp. n. Mesosoma with conspicuous silvery or golden tomentum on pronotal lobes, metanotum, sides of propodeum; gonostyles either angu- larly raised apicodorsally (fig. 51), or simply rounded" (fig. 4)" |e RR 12 Gonostyles simply rounded apicodorsally (fig. 54); tomentum of mesosoma golden or (So- lomon islands) pilosity brown; New Guinea, Solomon Islands ......... permutans (Turner) Gonostyles apicodorsally raised into a broad angle (fig. 51); tomentum of mesosoma silvery, pilosity pale; New Guinea, Bismarck IslandsYBiaksy tern egens (Kohl) Flagellomere III with small apical placoid, IV- VI with long, narrow, concave placoids (fig. 28); sternite V-VII with fringes of yellow set- osity; New.Guineab.r.: ein. ae 2 delicata sp. n. Flagellomere IV with long, narrow, concave placoid, V with small apical placoid (fig. 32); sternites without fringes, only with sparse yel- low pilosity; New Guinea ........ papua sp. n. Only flagellomeres IV-V with placoids (fig. 2) SEN VSN, PAPER ss cestra sp. n. Flagellomeres (II-) III-V with broad, convex Placoidi(tios 129433) Riss See 14 Flagellomere III 1.4 times as long as I (fig. 33); lower apical angle of gonostyle acute (fig. 50); facial tomentum usually silvery; Nepal to China and Singapore .......... diodon (Kohl) - Flagellomere III 1.1-1.2 times as long as I (fig. 29); lower apical angle of gonostyle a right angle (fig. 55); facial tomentum golden or pale golden; Sumatra, Borneo, Java, Nusa Tenggara, Moluccas, Philippines ........ severini (Kohl) Isodontia capillata sp. n. (figs. 26, 47) Type material. - Holotype @, Indonesia, Irian Jaya, Jayapura, xii.1936, W. Stüber (RMNH). — Paratypes: In- donesia: Irian Jaya, Jayapura, vii.1938, L. J. Toxopeus (1 6, RMNH). Papua New Guinea: Madang Province, Duru, 15 km SW. Madang, ca. 5° 20 S, 145° 43 E, 14.111.1987, W. J. Pulawski (1 8, CAS). Diagnosis Differing from all other species of this group by the dense yellow pilosity covering the posterior tergites, and by the shape of the gonostyles (fig. 47). Description Male. - Body length 15.0-16.5 mm, length of forewing 11.0-12.0 mm. Black; mesoscutum with fine oily shine; posterior margins of tergites broadly reddish to yellowish translucent. Wings light brownish hyaline, apically infuscated. Erect pilosity of head and mesosoma yellow; facial to- mentum golden; mesosoma with conspicuous golden tomentum on pronotum, sides of mesoscu- tum, metanotum, mesopleuron and sides of prop- odeum; metasoma with very conspicuous pale yel- low pilosity, dense and erect on posterior tergites. Clypeus 1.0 times as long as 10C, with vague median carina; IOV 1.1 times IOC; relative lengths of flagel- lomeres I-III 1: 1.4 : 2.6; third to sixth flagellomere with broad flat placoid. Mesoscutum densely rather finely punctate, punctures on disc 0.5-1 diameter apart; propodeal dorsum shiny, anteriorly trans- versely strigose, posteriorly very densely punctate. Petiole 1.2 times as long as hind basitarsus; ster- nites without fringes, only seventh sternite with some longer pilosity; eighth sternite apically trun- cate; gonostyle apically with strong tooth pointing upward. Etymology Capillata is an adjective, from Latin capillatus = long-haired. HENSEN: Malesian Sphecina Isodontia cestra sp. n. (figs. 7, 27, 48) Type material. - Holotype Q, ‘Indonesia, N. Sulawesi, 100 m, Tangkoko-Dua Saudara N. R., 1°30’ N, 125° 10’ E, 28.XI.1985, C. v. Achterberg, RMNH’86’ (RMNH). — Paratypes: Indonesia, Sulawesi: Dumoga-Bone N. P., ca. 220 m, nr. Base Camp Toraut R., 0°34’ N, 123°54 E, 10.xi.1985, C. v. Achterberg (1 9, RMNH); Kendari, iv.1874, O. Beccari (1 9, McG); North Sulawesi, Toli-Toli, xi-xii.1895, H. Fruhstorfer (2 4, IRSN, RMNH); 47 km WSW Kotamobagu, Dumoga Bone N. P., Toraut, 211 m, vi.1985, G. R. Else (2 9 4 6, BMNH, RMNH); Mt. Sapoetan, viii.1937, C. T. & B. B. Brues (1 9, MCz). Diagnosis Similar to severini and diodon; the female differs in having the petiole as long as hind basitarsus (longer in diodon and severini), the male has the placoids restricted to 4th and 5th flagellomere, and the gonostyles differ markedly in shape. Description Female. - Body length 15.5-18.5 mm, length of forewing 11.5-13.0 mm. Black; mesoscutum with oily shine; posterior margins of tergites brown. Wings yellowish, with sharply defined dark mark apically. Erect pilosity of head and mesosoma yel- low; facial tomentum golden; mesosoma with con- spicuous golden tomentum on pronotum, sides of mesoscutum, mesopleuron, metanotum, and sides and declivity of propodeum. Clypeus 1.1 times as long as IOC, without median carina; IOV 1.1 times 10C; relative lengths of flagellomeres I-III 1: 0.8 : 0.9. Mesoscutum densely rather coarsely punctate, punctures on disc 0.5-1 diameter apart; propodeal dorsum shiny, rather coarsely rugose-punctate. Pe- tiole 0.95-1.05 times as long as hind basitarsus. Male. - Body length 14.5-17.0 mm, length of forewing 10.5-12.0 mm. Like the female, except for the following. Erect pilosity of head and mesosoma white; facial tomentum silvery; mesosoma without conspicuous tomentum. ‘Clypeus with vague me- dian carina; IOV 1.2 times 10C; relative lengths of flagellomeres I-III 1: 1.0 : 1.15; fourth and fifth flagellomere with broad convex sensory area. Me- soscutum sparsely rather coarsely punctate, punc- tures on disc 1-2 diameters apart. Petiole 1.05 times as long as hind basitarsus; eighth sternite apically angularly emarginate; gonostyle in lateral view strongly constricted in the middle, apically swollen. Etymology Cestra is an arbitrary combination of letters, to be treated as a noun in apposition. 15 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 | Figs. 13-33. 13-18, Sphex, clypeus and mandibles. - 13, muticus, ®; 14, muticus, &; 15, solomon, 2; 16, solomon, 3; 17, praedator, &; 18, walshae, &. 19-33. Isodontia, proximal flagellomeres. - 19, aurifrons, 2; 20, pilipes, 9; 21, praslinia, 9; 22, aurifrons, &; 23, formosicola, &; 24, pilipes, 8; 25, chrysorrhoea, &; 26, capillata, 3; 27, cestra, 8; 28, delicata, 8; 29, diodon, 8; 30, egens, 6; 31, immaculata, 8; 32, papua, 8; 33, severini, &. 16 Isodontia delicata sp. n. (figs. 8, 28, 49) Type material. - Holotype 8, Indonesia, Irian Jaya, Jayapura, vii.1938, L. J. Toxopeus, Neth. Ind.-Amer. New Guinea Exp. 1939-39 (RMNH). — Paratypes: Indonesia: same data as holotype (5 6, RMNH, BMNH); same locality, vi.1937 (1 @), xii.1936 (2 &), W. Stüber (RMNH). Diagnosis Similar to papza (male, female in both species unknown), with which it shares the combination of extremely short flagellomeres I-III, and long nar- row concave placoid on fourth flagellomere; deli- cata however has similar placoids on fifth and sixth flagellomere, and fringes of yellow setosity on the posterior tergites; these traits are lacking in papa. Description Male. - Body length 13.5-15.5 mm, length of forewing 10.5-11.5 mm. Black; mesoscutum with oily shine; sides of first tergite and posterior mar- gins of other tergites and sternites reddish. Wings slightly yellowish hyaline, apically infuscated. Erect pilosity of head and mesosoma yellow; facial to- mentum golden; mesosoma with conspicuous golden tomentum on pronotum, sides of mesoscu- tum, mesopleuron, sides of propodeum; fourth to eighth sternite with fringes of yellow setosity. Cly- peus 1.0 times as long as IOC, with vague median carina; IOV 1.0 times IOC; relative lengths of flagel- lomeres I-III 1: 1.5 : 2.1; third flagellomere with small apical placoid, fourth to sixth with narrow concave placoid. Mesoscutum densely rather coar- sely punctate, punctures on disc 0.5-1 diameter apart; propodeal dorsum shiny, anteriorly trans- versely strigose, posteriorly punctate-strigose. Pe- tiole 1.2-1.3 times as long as hind basitarsus; eighth sternite truncate; gonostyle simple. Etymology Delicata is an adjective, from Latin delicatus = delicate. Isodontia diodon (Kohl) (figs. 29, 50) Sphex (Isodontia) diodon Kohl, 1890: 377. Lectotype & (present designation): ‘Celebes, Sumatra’ (NMW!!) Sphex maia Bingham, 1894: 379. Syntypes, 6, 9: Burma, Tenasserim (BMNH!!). Distribution. - This species occurs on the Asian continent, from Nepal to China and Peninsular Malaysia. Notes. - 1. The lectotype of Sphex diodon Kohl is a male, labelled ‘Stevens, 860’, ‘diodon typ, det Kohl’ (NMw). Kohl gave as the habitat of the spe- HENSEN: Malestan Sphecina cies ‘Celebes, Sumatra’, but this is almost certainly incorrect. The species that corresponds to both the type and the description is found only on the Asian continent. 2. I. diodon has been regarded as a polytypic species by past workers (Van der Vecht 1957, Bo- hart & Menke 1976). However, typical I. diodon differs in several morphological aspects from the forms associated with it in the past. I. nigelloides (Strand) is a distinct species. Three other forms are treated here as belonging to I. severini (Kohl). The three species are allopatric: I. nigelloides [stat. n.) is restricted to Southern India and Sri Lanka; I. diodon occurs on the Asiatic mainland from Nepal to China and Malaysia, whereas I. severini occurs in the Archipelago. Isodontia egens (Kohl) (figs. 30, 51) Sphex (Isodontia) egens Kohl, 1898: 335. Holotype &: New Britain, Kinigunang (TMB!!). Distribution. - New Britain, Western New Gui- nea, Biak. Isodontia franzi (Cameron) Sphex (Isodontia) franzi Cameron, 1902: 246. Holotype 4: Borneo (BMNH!!). Distribution. - Only the type specimen from Borneo is known. Isodontia immaculata sp. n. (figs. 31, 52) Type material. - Holotype d, Papua New Guinea, New Ireland, Trail to Lelet, Plateau near Danum, 15.xii.1969, J. E. Tobler (CAs). - Paratypes: Papua New Guinea: same data as holotype (2 6, CAS, RMNH); Lavongai [New Han- over], Banatam, 24.iii.1962, Noona Dan Exp. (1 9, UZMC). Diagnosis Within the diodon-group, this species is the least tomentose one: the facial tomentum is very sparse and silvery, the mesosoma lacks tomentum, whereas all other species have conspicuous spots of silvery or golden tomentum on pronotal lobes, angles of propodeum, etc. Structurally, the spe- cies is similar to egens and permutans, males of these species share the combination of extremely short proximal flagellomeres, and broad flat pla- coids on third to sixth flagellomere; immaculata differs markedly in the shape of the male gono- style. 197 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Description Female. - Body length 17.5 mm, length of fore- wing 12.5 mm. Black. Wings clear, forewing ante- riorly and apically infuscate, the dark mark not sharply delimited. Erect pilosity of head and mes- osoma white; facial tomentum silvery, sparse; mes- osoma without tomentum. Clypeus 1.1 times as long as IOC, without median carina; IOV 1.0 times 10C; relative lengths of flagellomeres I-III 1: 0.95 : 1.0. Mesoscutum densely rather coarsely punctate, punctures on disc 0.5-1 diameter apart; propodeal dorsum shiny, rather coarsely rugose-punctate. Pe- tiole 1.1 times as long as hind basitarsus. Male. - Body length 16.0 mm, length of forewing 11.5 mm. Like the female except for the following. Clypeus 1.1 times as long as IOC, with vague median carina; relative lengths of flagellomeres I-III 1 : 1.4 : 2.4 ; third to sixth flagellomere with broad flat sensory area. Mesoscutum densely rather coarsely punctate, punctures on disc 1-1.5 diameters apart; propodeal dorsum shiny, rather coarsely rugose-punctate. Pe- tiole 1.1 times as long as hind basitarsus; eighth sternite apically not emarginate; gonostyle apico- dorsally swollen and rounded. Etymology Immaculata is an adjective, from Latin zmmacu- latus = immaculate. Isodontia papua sp. n. (figs. 32, 53) Type material. - Holotype @, Indonesia, New Guinea, ‘Neth. Ind.-American New Guinea Exped., Araucaria Camp, 800 m, 30.III.1939, L. J. Toxopeus’ (RMNH). Diagnosis Similar to delicata (male, female of both species unknown), with which it shares the combination of extremely short proximal flagellomeres, and a long narrow, concave placoid on the fourth flagelio- mere. However, delicata has similar placoids on fifth and sixth flagellomere, whereas papua only has a vestigial one on fifth, and none on sixth. Additionally, papua lacks fringes of yellow setosity on the sternites, as found in delicata. Description Male. - Body length 15.0 mm, length of forewing 11.5 mm. Black; mesoscutum with oily shine; pos- terior margins of tergites and sternites reddish to yellow. Wings light brownish hyaline, apically in- fuscated. Erect pilosity of head and mesosoma yel- low; facial tomentum silvery; mesosoma with con- spicuous golden tomentum on pronotum, sides of mesoscutum, mesopleuron, etc.; posterior tergites 18 with appressed pale yellow pilosity. Clypeus 1.0 times as long as IOC, with vague median carina; IOV 1.1 times IOC; relative lengths of flagellomeres I- II 1: 1.45 : 2.0; fourth flagellomere with long, narrow, concave placoid, fifth with small apical placoid. Mesoscutum densely rather coarsely punc- tate, punctures on disc 0.5-1 diameter apart; prop- odeal dorsum shiny, anteriorly transversely punc- tate-strigose, posteriorly reticulate-punctate. Petiole 1.2 times as long as hind basitarsus; sixth to eighth sternite with scattered pilosity poste- riorly, without real fringes; eighth sternite trun- cate; gonostyle simple. Etymology Papua is noun in apposition, it is the common name for the native inhabitants of New Guinea. Isodontia permutans (Turner) (fig. 54) Sphex permutans Turner, 1912: 198. Lectotype 9 (pres- ent designation): New Guinea, Wataikwa River (BMNH!!) Distribution. - New Guinea, Solomon Islands. Notes. - 1. The males of permutans and its closest relatives are all easily recognized using characters of the genitalia and secondary sexual characters. The females of permutans, papua, del- icata, and capillata are apparently inseparable. As the type of permutans is a female, the identity of this species is uncertain. I have associated the most common and widely distributed type of male with permutans, and described the others as new spe- cies. I have not been able to associate females with these. If characters are eventually found to identify females of these species, my interpretations may need revision. 2. I. permutans is geographically variable with respect to the colour of pilosity and tomentum. Typically (New Guinea) the pilosity is pale yellow, the tomentum golden. Specimens from Bougain- ville are similar to the typical form. The pilosity of head and mesosomal dorsum is partly to entirely dark in specimens from the other Solomon Islands. The tomentum is silvery in specimens from Gizo, Kolombangara, San Cristobal, and Malaita. Isodontia severini (Kohl) stat. rev. (figss 3325) Sphex (Isodontia) severini Kohl, 1898: 337, 9. - Java (IRSN). Sphex malayanus Cameron, 1902: 134. Lectotype & (present designation): Borneo, Kuching (BMNH!!) Sphex (Isodontia) severini var. philippensis Rohwer, 1921: 669. Holotype 9: Palawan (USNM!!). Isodontia diodon alemon Vecht, 1957: 367. 4: Sumba (NMB) Syn. n. Distribution. — Malaysia, Singapore, Indonesia (Sumatra, Java, Borneo, Sumba, Halmaheira), Phi- lippines. Group of Isodontia ochroptera Diagnosis Body length ¢ 16-22 mm, 9 20-22 mm. Labrum of female with median carina; mandible in both sexes tridentate, basal inner tooth in male strongly developed, projecting beyond second inner tooth; clypeus of female with pair of apical teeth medially; first flagellomere in female distinctly longer than second, in male first and second flagellomere equally long; third to eighth flagellomere of male with broad, convex, papillate sensory areas; an- tenna of female without sensory areas; outer side of hind tibia with five spines; petiole a little shorter than hind basitarsus. Species included Species treated. - I. ochroptera (Kohl). Other included species. — I. sonani (Tsuneki) from Taiwan. Isodontia ochroptera (Kohl) (figs. 9-10, 34) Sphex (Isodontia) ochropterus Kohl, 1890: 378. 2: In- donesia, Sulawesi, Bantimurung (Mus. Dresden, prob- ably destroyed). Distribution. - Continental Asia from Fastern India to Laos and Malaysia, Indonesia (Sumatra, Borneo, Java, Sulawesi, Aru, Halmaheira), Philip- pines (Palawan, Samar) Group of Isodontia praslinia Diagnosis Body length & 15-23 mm, 9 16-23 mm. Labrum of female with a pair of short, parallel, projecting carinae apically; mandible of both sexes tridentate, basal inner tooth strongly developed, projecting beyond second inner tooth; clypeus of female with median notch; first flagellomere longer than se- cond; flagellum of male with generally with flat placoids on flagellomeres IV-VI; antenna of female without sensory areas. Scutellum and metanotum convex; propodeal dorsum more or less reticulate; outer side of hind tibia with 5-10 spines. Petiole usually shorter than hind basitarsus. HENSEN: Malesian Sphecina Species included Species treated. - I. e/sez sp. n., jaculator (Smith), nidulans sp. n., praslinia (Guérin-Méneville), se- picola (Smith). Other included species. - I. obscurella (Smith) from Australia, I. xanthognatha (Perez) (= nigella (Smith), nec Lichtenstein) [stat. n.] from contin- ental Asia. Key to species 1. Pilosity of head and mesosoma dark brown to black; wings dark brown with violaceous shine; Timor (?), New Guinea and neighbouring is- la ads praslinia (Guérin-Méneville) — Erect pilosity of head and mesosoma white to yellowish; colour of wings variable .......... 2 2. Propodeal dorsum sparsely punctate, interspa- ces smooth, shiny; interocular distance across vertex shorter than across clypeus; Philippines ren. sr cht dal nidulans sp. n. - Propodeal dorsum punctate-reticulate to stri- gose-punctate, more or less dull, without SMOothinters paces ks. "dr Cee 3 3. Facial tomentum sparse, silvery, sculpture of clypeus easily visible; Moluccas ................. RE RR OA jaculator (Smith) - Facial tomentum dense, golden or silvery, sculpture of clypeus largely obscured ........ 4 4. Petiole of female 0.8-0.95, of male 0.9-1.0 times as long as hind basitarsus; IOV 0.85-1.05 times IOC; male: mandibles entirely bright red- dish; New Guinea and neighbouring islands, Northern Australia .......... sepicola (Smith) — Petiole of female 0.95, of male 1.05-1.1 times as long as hind basitarsus; IOV 1.1 times IOC; male: basal third of mandibles black; Sulawesi reen teater ae cai elsei sp. n. Isodontia elsei sp. n. (fig. 35) Type material. - Holotype 8, Indonesia, Sulawesi, ‘Ro- senberg, Toelabella, Celebes’ (RMNH). — Paratypes. In- donesia, Sulawesi: Gorontalo (1 &), Kwadang (1 @), both Rosenberg (RMNH); Patunuang, i.1896, H. Friihstorfer (2 Q, RMNH); 47 km WSW Kotamobagu, Dumoga Bone N. Pk., Toraut (Forest Edge), 211 m, v-vi.1985, G. R. Else (6 3, 3 9, BMNH, RMNH); Luwuk, Matanyo Forest, N. of Kayutano, 120 m, 20.x.1989, C. van Achterberg & M. Tulung (1 9, RMNH). Diagnosis Very similar to sepicola, from which it differs only in the characters mentioned in the key. Description Female. - Body length 19.0-20.5 mm; length of forewing 13.5-15.5 mm. Black; mesoscutum with 19 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 oily shine. Wings light brownish, apical margins more strongly infuscated, with violaceous shine. Erect pilosity of head and mesosoma nearly white; facial tomentum golden, dense, leaving sculpture of clypeus visible under a certain angle only; meso- soma without conspicuous tomentum. Clypeus 0.8 times as long as 10C, without discernable median carina; IOV 1.1 times 10C; relative length of flagel- lomeres I-III 1 : 0.8 : 0.8. Mesoscutum shiny, spar- sely finely punctate, punctures on disc 1-3 diame- ters apart; propodeal dorsum more shiny than in I. sepicola, weakly transversely rugose and shal- lowly punctate. Petiole 0.95 times as long as hind basitarsus. Male. - Body length 15.5-20.5 mm; length of forewing 13.5-15.5 mm. Like the female except for the following. Black; mandibles with short reddish zone just before apex. Wings brownish hyaline, apical margins infuscated. Erect pilosity of head and mesosoma yellowish to white. Clypeus 0.85 times as long as 10C; relative length of flagello- meres I-III 1 : 0.8 : 0.85. Petiole 1.05-1.1 times as long as hind basitarsus; third to seventh sternite with fringes of brown setosity; gonostyle simple. Etymology Elsei is a noun in the genitive case, referring to Mr. George R. Else, collector of most of the type series, and many other interesting Sulawesi wasps. Isodontia jaculator (Smith) (fig. 36) Sphex jaculator Smith, 1860: 122. Holotype &: Indonesia, Bacan (OUM!!). Distribution. — Exclusively known from the is- land Bacan, in the Indonesian Northern Moluccas. Isodontia nidulans sp. n. Type material. - Holotype 9, Philippines, Luzon, ‘Olongapa, P. I., Subic Bay, V.1967’, ‘Coll. by J. C. Thom- son’ (USNM). Diagnosis Immediately separated from other species of this group by its very fine sculpture; in particular the propodeal dorsum is smooth, shiny, with sparse fine punctuation. Description Female. - Body length 20.0 mm; length of fore- wing 15.5 mm. Black; mesoscutum with oily shine. Wings brown, with violaceous shine. Erect pilosity of head and mesosoma white; facial tomentum silvery, sparse; mesosoma without conspicuous to- 20 mentum. Clypeus 0.7 times as long as 10C, with blunt median carina on dorsal half; 10v 0.95 times 10C; relative length of flagellomeres I-II 1 : 0.75 : 0.75. Mesoscutum shiny, obsoletely punctate; prop- odeal dorsum shiny, along anterior margin with weak rugae, remaining part smooth, finely, shal- lowly punctate. Petiole 0.9 times as long as hind basitarsus. Etymology Nidulans is an adjective based on the Latin verb nidulare = to shine. Isodontia praslinia (Guérin-Méneville) (figs. 11-12, 21, 37) Sphex praslinius Guérin-Méneville, 183 1: pl. 8. Holotype Q: New Ireland, Pt. Praslin (MCG!). Sphex (Isodontia) ustulata Kohl, 1890: 378. Holotype 9: Timor (NMW!!). Syn. n. Sphex (Isodontia) simplex Kohl, 1898: 334. Lectotype & (present designation): New Guinea: Erima, Astro- labe-Bay (TMB!!). Syn. n. Sphex (Isodontia) insularis Cameron, 1901: 240. Holo- type 6: New Britain (BMNH!!). Syn. n. Distribution. - This species occurs on New Gui- nea and neighbouring islands: New Britain, New Ireland, Misool Island, the Solomon Islands. The record from Timor is based only on the type of ustulata, and it is doubtful whether the species indeed occurs there. Species whose occurrence is centered on the Papuan subregion, like I. praslinia, and also for example I. sepicola and S. cognatus rarely occur in the Lesser Sunda islands. The two areas are climatically quite different. Isodontia sepicola (Smith) (fig. 38) Sphex sepicola Smith, 1859: 158. Lectotype 9 (present designation): Aru (OUM!!) Sphex albohirtus Turner, 1908: 466. Lectotype (present designation): Australia: Mackay (BMNH!!). Syn. n. Distribution. - This species occurs on New Gui- nea, and neighbouring islands: Aru islands, So- lomon islands, Misoöl, and in Australia, where it seems to be restricted to Northern Queensland. Genus Sphex Linnaeus Group of Sphex argentatus Diagnosis Body length & 20-28 mm, 9 23-34 mm. Mand- ible of both female and male bidentate; antenna of male with placoid(s), at least on fifth flagellomere; scutellum convex, medially impressed; metanotum medially raised into a pair of tubercles, or into a single medially impressed tubercle; claw teeth large, at sharp angle to claw. Species included Species treated. — S. alacer Kohl, argentatus Fa- bricius, diabolicus Smith, finschu Kohl, madasum- mae Van der Vecht, sericeus Fabricius Other included species. — carbonicolor Vecht, decoratus Smith, ephippium Smith, modestus Smith (all Australian), fumicatus Christ (Africa to SW. India and Sri Lanka). Key to species 1. Propodeal dorsum with 4-6 transverse ridges - Propodeal dorsum without ridges ........ pig 2. Female: apical margin of clypeus with median tooth; male: 2nd to 6th flagellomere with broad placoid (fig. 39); New Guinea............ BANE OE SR dara alacer Kohl - Female: apical margin of clypeus without tooth; male: 4rd or 4th to 6th flagellomere with placoid (fig. 42); Sri Lanka to New Guinea bbl ark aisha ee sericeus (Fabricius) 3. Wings uniformly dark brown, with violaceous effulgence; Burma to New Guinea, Philippines A bia teeta melas. madasummae Van der Vecht - Wings clear or yellowish, only at basis and apex sometimessdarkenede eh no 4 4. Wings strongly yellowish to yellow-brown; Sri lanka to Sulawesi ............ diabolicus Smith - Wings essentially clear, infuscate at base and Aperen kend Mantes ile. 5 5. Metanotum with single medially impressed tubercle; male: 4th to 6th flagellomere with broad placoids (fig. 41); Moluccas, New Gui- nea, Solomon Islands, Australia finschi Kohl — Metanotum strongly bituberculate; male: only Sth flagellomere with narrow placoid (fig. 40); widespreade nn argentatus Fabricius Sphex alacer Kohl (fig. 39) Sphex alacer Kohl, 1895: 54. Holotype 9: New Guinea (NMW!). Distribution. - New Guinea, New Ireland. Sphex argentatus Fabricius (fig. 40) Sphex argentatus Fabricius, 1787: 274. Lectotype 9 (de- signated by Van der Vecht 1961: 28): India, Coroman- del (Mus. Lund!). HENSEN: Malesian Sphecina Sphex umbrosus Christ, 1791: 293, pl. 29. No locality given [type lost]. Sphex argenteus Turton, 1800: 485. India, Coromandel [possible emendation]. Sphex argentifrons Lepeletier, 1845: 337. Lectotype à (designated by Menke in Bohart & Menke 1975: 114): Java (Mus. Torino!). Sphex plumifer Costa, 1864: 112. Philippines, Luzon (Mus. Naples). Sphex umbrosus var. nanulus Strand, 1913: 83. Holotype Q: Taiwan, Taihorin (DEI!!). Sphex umbrosus var. fumosus Kohl, 1890: 408. Japan (TMB). Distribution. - Sri Lanka and India to Indochina and Japan; Malaysia, Indonesia, Philippines, Papua New Guinea and NE. Australia. Note. - Sphex unicolor Fabricius, 1787, listed as a synonym of argentatus by Bohart & Menke (1975), was described from Spain where argentatus does not occur. Van der Vecht (1961) suggested that the name might refer to S. fumicatus Christ, 1791, but that species has never been reported from Spain either. The only Spanish species of Sphex that agrees with the description is S. afer Lepeletier. Alternatively, the locality may be incor- rect. Sphex diabolicus Smith Sphex diabolicus Smith, 1858: 100. Lectotype 9 (desig- nated by Van der Vecht 1973): Sarawak (OUM!) Sphex flammitrichus Strand, 1913: 83. Lectotype 2 (pres- ent designation): Taiwan (DEI!!) Sphex umbrosus var. aureopilosus Berland, 1928: 330. Lectotype @ (designated by Menke in Bohart & Menke 1975): Ba-Cha, Tonkin (MNHN!!) Sphex fulvohirtus Bingham, 1890: 242. Holotype 9: Sri Lanka, Pundalaya (BMNH!!). Syn. n. Types. - The lectotype of Sphex flammitrichus is a female labelled ‘Kankau (Koshun), Formosa, H. Sauter, vii.1912’, the paralectotypes are 3 males, labelled ‘Korosan, Formosa, Sauter, xi.07’ (all DEI). Distribution. - India and Sri Lanka to Malaysia, Indonesia (Sumatra and neighbouring islands, W. Java, Borneo, Sulawesi), Philippines. Sphex finschii Kohl (fig. 41) Sphex finschu Kohl, 1890: 412. Lectotype & (present designation): New Britain (ZMB!!) Distribution. - Indonesia (Ambon, Aru Islands, Irian Jaya), Papua New Guinea, Bismarck Islands, Solomon Islands, Australia. 21 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Sphex madasummae Van der Vecht Sphex caerulescens Guillou, 1841: 320. Holotype £: Bor- neo (MNHN!!) [non Reich, 1795]. Sphex maurus Smith, 1856: 255. Syntypes, 9, &: Sulawesi (BMNH!!) [non Fabricius, 1787]. Sphex nigerrimus Costa, 1864: 112. Philippines, Luzon (Naples Museum?) [non Scopoli, 1763]. Sphex madasummae Van der Vecht, 1973: 345 [new name for Sphex maurus Smith]. Distribution. - Malaysia, Indonesia (not known from Nusa Tenggara); Philippines, Papua New Guinea. Sphex sericeus Fabricius (fig. 42) Sphex aurulentus Fabricius, 1793: 201. - India, Tranqu- ebar (location of type unknown) [non Fabricius, 1787]. Sphex sericeus Fabricius, 1804: 211. Lectotype 9 (desig- nated by Van der Vecht 1961): ‘in maris pacifici Insu- lis’ (Mus. Lund!) Sphex fabricu Dahlbom, 1843: 27. 9: India, Tranquebar (location of type unknown). Sphex ferrugineus Lepeletier, 1845: 345. 9: ‘sans patrie, mais exotique’ (MNHN?). Sphex lineolus Lepeletier, 1845: 353. &: Java (MNHN?) [locality probably erroneous]. Sphex ferox Smith, 1862: 55. Syntypes, 9, 4: Ambon, Sulawesi (BMNH!!, OUM) [non Westwood, 1837]. Sphex lepeletierii Saussure, 1867: 40. Holotype 9: Java (NMW!!) [locality probably erroneous |. Sphex godeffroyi Saussure, 1869: 57. Q: Australia, Queensland, Cape York (Mus. Hamburg). Sphex aurifex Smith, 1873: 460, pl. xliv, fig. 3. Holotype Q: NW. Australia, Champion Bay (BMNH!!). Sphex aurulentus var. pallidehirtus Kohl, 1890: 393. Syn- types 6: New Guinea: Port Moresby; Ambon (NMW?). Sphex rugosus Matsumura, 1912: 176, 53, fig. 8. 9: Tai- wan: Horisha, Koshun. Sphex sericeus ferocior Van der Vecht & Krombein, 1955: 40 [new name for Sphex ferox Smith]. Sphex sericeus nigrescens Van der Vecht & Krombein, 1955: 39. Holotype ©: Philippines, Luzon, Los Bafios (USNM). Sphex sericeus stueberi Van der Vecht & Krombein, 1955: 42. Holotype Q: Indonesia, Irian Jaya, Jayapura (RMNH!!). Sphex sericeus wegneri Van der Vecht & Krombein, 1955: 39. Holotype &: Indonesia, Kalimantan, Sama- rinda, Mura Kaman (RMNH!!). Distribution. - India and Sri Lanka to China and Indochina, Malaysia, Indonesia, Philippines, Papua New Guinea, N. Australia. Note. - This species occurs in many colour- forms, which are largely allopatric. Van der Vecht and Krombein (1955) recognized eight subspe- cies. 22 Group of Sphex resplendens Smith Diagnosis Body length & 17-24 mm, 9 20-33 mm. Mand- ible of both female and male bidentate; antenna of male without placoids; scutellum of female flat or very weakly convex, conspicuously shiny, of male convex, medially not or hardly impressed; claw teeth small, perpendicular to claw. Species included Species treated. - S. brachystomus Kohl, con- frater Kohl, dorycus Guérin-Méneville, mimulus Turner, muticus Kohl, resplendens Kohl, solomon sp. n., wilsoni sp. n. Other included species. - S. fumipennis Smith, S. gilberti Turner, S. /uctuosus Smith (all from Aus- tralia), and S. antennatus Smith stat. n. (from New Caledonia, New Hebrides and Loyalty Islands; listed as a subspecies of S. fumipennis by Bohart & Menke 1976). Key to species 1. Metasoma partly red (sometimes only part of POSTEIOLICELOILES) REI eee PAPER eee a 2 = 'Metasoma entirely black ist. 4 2. Pilosity of mesosoma white; 8th sternite of male with narrow apical process; Australia, News Guinea AR mimulus Turner - Pilosity of mesosoma dorsally yellow to black, ventrally brownish to black; 8th sternite of male triangular, without process ............ 3 3. Metasoma except petiole entirely red; pilosity of mesosoma dorsally yellow to brownish; pe- tiole of female longer than wide; gonostyles of male without inner lobe; New Guinea DEE, AMI MOREE oe dorycus Guérin-Méneville — Metasoma with only posterior tergites and sternites red; pilosity of mesosoma black; pe- tiole of female as long as wide; gonostyles of male ventrally with flat, almost circular inner lobe (fig. 56); Solomon and Bismarck Islands TUE LEMON O sil rara confrater Kohl 4. Mesonotum densely covered with silvery or golden tomentum, completely obscuring sculp- WILLIE 5 - Mesonotum without or with sparse tomentum, sculpturesvisibleichroushout ame 6 5. Tomentum golden; wings yellowish, apical margins infuscate; New Guinea ................ ACRI ER OLE wilsoni sp. n. - Tomentum silvery; wings dark brown, with violaceous effulgence; New Guinea, Bismarck Islands on RETE brachystomus Kohl 6." Bemalesıı 1. Mes TALL ed ren 7 MOMENT E NEN a EEE: ©) 7. Mandibles long, strongly curved (fig. 13), when closed reaching beyond base of opposite mandible; apical margin of clypeus strongly arcuate; median carina of labrum projecting into a small apical tooth; Buru, Ambon BEER So tri eG I muticus Kohl - Mandibles short, moderately curved (fig. 15), when closed not reaching base of opposite mandible; anterior margin of clypeus weakly arcuate; median carina of labrum not project- Inga picallll Ake Australian di ae ithe, Muineent; 8 8. Interocular distance at vertex longer than combined lengths of flagellomeres II + III; metanotum without silvery tomentum; Moluc- cas, New Guinea, Philippines, Australia MERI Rn resplendens Kohl — Interocular distance at vertex equal to com- bined lengths of flagellomeres II + III; meta- notum densely covered with silvery tomentum; Solomon Islands ................ solomon sp. n. 9. Mandibles with basal half straight, sharply curved inward distally (fig. 14); petiole in dor- sal view more than 2 times as long as wide; gonostyles not emarginate, the apex simple; BUGURATIDONERE SR muticus Kohl — Mandibles smoothly curved, approximately a quarter-circle (fig. 16); petiole in dorsal view less than 2 times as long as wide; gonostyles dorsally emarginate, the apex produced into tooth pointing upward (fig. 58) ............ 10 10. Pilosity of head and mesosoma dark brown to black; wings dark brown; Philippines, Moluc- caseiNewGuineasAustraliad SER CNR Ans esi vio EK Din resplendens Kohl - Pilosity of head and mesosoma yellowish, partly dark brown on head and mesosomal dorsum; wings brownish hyaline; Solomon Is- lands. esi esi pal Dal solomon sp. n. Sphex brachystomus Kohl Sphex brachystomus Kohl, 1890: 415. Syntypes 9 4: New Britain (NMW!). Distribution. - Papua New Guinea, Bismarck Islands (New Ireland, New Britain). Sphex confrater Kohl (fig. 56) Sphex confrater Kohl, 1890: 414. Syntypes 9: New Bri- tain (Mus. Hammon’). ? Sphex confrater var. sieberti Strand, 1910: 46. Syntypes Q: Java, Batavia (location of types unknown). Distribution. - Solomon Islands (Kolomban- gara, Guadalcanal), Papua New Guinea, Bismarck Islands (New Britain, New Ireland). HENSEN: Malesian Sphecina Note. — Judging on the type-locality, it seems very unlikely that steberti Strand is indeed a variety of confrater, but it is not clear from the description to which other species it might pertain. Sphex dorycus Guérin-Méneville Sphex dorycus Guérin-Méneville, 1838: 262. Holotype 9: New Guinea, Manokwari (‘Dory’) (MCG). Sphex errabundus Kohl, 1898: 338. Holotype 2: Indone- sia, Waigeo ( Waigiu') (TMB!). Distribution. - Indonesia (Waigeo, Irian Jaya), Papua New Guinea. Note. - The synonymy of this species was dis- cussed by Van der Vecht (1973). Sphex mimulus Turner Sphex mimulus Turner, 1910b: 419. Holotype 9: Austra- lia, Queensland, Cairns (BMNH!). Distribution. - Australia (Queensland), S. Papua New Guinea. Sphex muticus Kohl (figs. 13-14, 57) Sphex muticus Kohl, 1885: 199. Lectotype @ (designated by Van der Vecht 1973): Indonesia, Ambon (NMW!) Distribution. - Indonesia (Buru, Ambon). Sphex resplendens Kohl Sphex nitidiventris Smith, 1859: 158. Holotype 9: In- donesia, Buru (OUM!) [non Spinola, 1851]. Sphex gratiosus Smith, 1859: 158. Holotype @: Indone- sia, Aru Islands (OUM!) [non Smith, 1856]. Sphex resplendens Kohl, 1885: 200 [new name for S. nitidiventris Smith]. Sphex gratiosissimus Dalla Torre, 1897: 424 [new name for S. gratiosus Smith, 1859]. Sphex wallacei Turner, 1908: 467 [new name for S. nit- tdiventris Smith]. Sphex lanceiventris Vachal, 1908: 23 [new name for S. nitidiventris Smith]. Sphex mertoni Strand, 1911: 231. Holotype 2: Indonesia, Aru Islands, Kobroör, Seltutti (Mus. Senckenberg!!). Distribution. - Philippines (Negros, Luzon); In- donesia (Aru Islands, Key Islands, Bacan, Ambon, Irian Jaya); Papua New Guinea, Australia (Queens- land). A few specimens from Bangka in the RMNH are probably incorrectly labelled. 25 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 34-61. Isodontia and Sphex. 34-45, proximal flagellomeres of 8. - 34, I. ochroptera; 35, I. elset, 36, I. jaculator, 37,1. praslinia; 38, I. sepicola; 39, S. alacer; 40, S. argentatus; 41, S. finscht, 42, S. sericeus; 43, S. rex; 44, S. subtruncatus, 45, S. walshae. 46-58. Gonostyle of 4, 46-55 + 57-58, lateral view, 56, ventral view. — 46, 1. pilipes; 47, I. capillata; 48, I. cestra; 49, I. delicata; 50, I. diodon; 51, L egens; 52, I. immaculata; 53, I. papua; 54, I. permutans; 55, I. severint, 56, S. confrater; 57, S. muticus; 58, S. solomon. 59-60, tarsal claw. 59, S. solomon, Q; 60, S. subtruncatus, Q. 61, S. rex, G, eighth sternite. 24 Sphex solomon sp. n. (figs. 15-16, 58-59) Type material. - Holotype 4, Solomon Islands, Guadal- canal Is., Lunga, 2.xii.1934, H. T. Pagden (RMNH). — Para- types: Solomon Islands: Tulagi (3 2), Guadalcanal, Lunga (1 Q 1 8), Ganonga (1 9 5 8), Nygela (1 ©), all H. T. Pagden (RMNH); Guadalcanal, Kukum, P. Greenslade (1 ©, BMNH); Isabel, Tatamba (1 9, Australian National Insect Collection, Canberra; 1 9, RMNH); Bougainville, Buin, G. Daniels (1 9, Australian Museum, Sydney); Gua- dalcanal, Lunga River Bridge, J. Schenk (1 &, Los Angeles County Museum); Guadalcanal, Kukum, E. S. Brown (1 & 7 9, BMNH, 2 9, RMNH). Diagnosis Similar to resplendens and brachystomus, with which solomon shares relatively short and weakly curved mandibles, and an apical lobe on the male gonostyle. It differs from these species in vestiture: the silvery tomentum of the mesosoma is restricted to the pro- and metanotum, particularly conspicu- ous on the latter, in male, the pilosity of head and mesosoma is basically pale. Structural features are: in female, lov equals flagellomere 2 + 3 (longer in resplendens and brachystomus); in male the apical lobe of the gonostyle is broad and rounded (narrow, pointed in the other species). Description Female. - Body length 20-25 mm; length of forewing 15.0-18.5 mm. Black, metasoma with fine greenish metallic shine. Wings brown. Pilosity of head and mesosoma dark brown; facial tomentum brownish to silvery; mesosoma with conspicuous silvery tomentum on metanotum and sometimes on pronotum. Clypeus 0.8 times as long as IOC, with weak median keel on dorsal half, apical margin slightly raised, with a pair of short teeth medially; IOV 0.75 times IOC; labrum with median carina, which does not project apically; relative lengths of flagellomeres I-III 1 : 0.6 : 0.6. Mesopleuron spar- sely finely punctate, interspaces punctulate, shiny; mesoscutum densely, rather coarsely punctate, punctures 0.5-1 diameter apart, interspaces smooth, shiny; scutellum slightly convex, sparsely finely punctate, shiny; propodeum coriaceous, dull. Fore basitarsus with 8 rake spines, median spines 2 time tarsal width, apical spine as long as second tarsomere. Petiole 0.3 times as long as hind basi- tarsus, in dorsal view hardly longer than wide. Male. - Body length 19-24 mm; length of fore- wing 13.5-17.0 mm. Like the female except for the following. Wings brownish hyaline, apical margins infuscate. Facial tomentum silvery; pilosity of head and mesosoma basically pale, more or less mixed with dark hairs or entirely dark on vertex, temples and mesosomal dorsum. Clypeus 0.95 times as long HENSEN: Malesian Sphecina as IOC, apical margin untoothed, medially de- pressed; IOV 0.85 times IOC; labrum ecarinate; scu- tellum convex, faintly impressed along midline. Apical margins of second to seventh sternite with row of black setae; eighth sternite apically pro- duced into a tooth, distal third medially keeled; gonostyle apically with short lobe pointing up- ward, ventrally in the middle with almost circular lobe pointing inward. Etymology Solomon is a noun in apposition, referring to the type locality. Sphex wilsoni sp. n. Type material. - Holotype 9, Papua New Guinea, Huon Peninsula, Mongi-Mape Watersheds, Nganduo to Yunzain, 1000-1500 m, 56.iv.1955, E. O. Wilson (MCz). Diagnosis Immediately separated from other species of the resplendens-group by the conspicuous golden to- mentum on head and thorax, and the yellow wings. Structurally, wilsont is similar to confrater, mimu- lus, dorycus and muticus, with which it shares the long, strongly curved mandibles. Description Female. - Body length 25 mm; length of fore- wing 19 mm. Black; metasoma with fine metal- lic shine. Wings yellow, apical margins infuscate. Pilosity of head and mesosoma dark brown; head, pro-, meso- and metanotum with dense golden tomentum, completely obscuring the sculpture of these parts. Clypeus 0.75 times as long as IOC, with weak median carina dorsally; apical margin slightly raised, with a pair of short teeth medially; 10v 0.75 times 10C; labrum with median carina, projecting apically into a short tooth; mandibles long, strongly curved, when closed reaching beyond each other’s bases (cf. fig. 13); relative length of flagel- lomeres I-III 1 : 0.6 : 0.65. Mesopleuron sparsely finely punctate, interspaces weakly punctulate; me- sonotum apparently (obscured by vestiture) den- sely finely punctate, shiny; propodeum finely cori- aceous, dull; fore basitarsus with 9 rake spines, median spines 1.4 times tarsal width, apical spine shorter than second tarsomere. Petiole 0.25 times as long as hind basitarsus, in dorsal view hardly longer than wide. Etymology Wilson: is a noun in the genitive case, referring to E.O. Wilson, collector of the only known spec- imen of this species. 25 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Group of Sphex subtruncatus Dahlbom Diagnosis Body length & 16-28 mm, Q 15-36 mm. Mand- ible of both male and female bidentate; antenna of male with placoid(s); scutellum convex, medially impressed; metanotum simple; claw teeth large, at sharp angle to claw. Species included Species treated. - S. cognatus Smith, formosellus Van der Vecht, praedator Smith, rex sp. n., staudın- geri Gribodo, subtruncatus Dahlbom, walshae sp. n. Other included species. - S. obscurus (Fabricius) (Sri Lanka to Thailand, ahasverus Kohl, semifos- sulatus Van der Vecht, vestitus Smith, ermineus Kohl, and bilobatus Kohl (all Australian). Note. - This is obviously a heterogeneous as- semblage. The species are grouped together only on basis of the lack of specialized features defining the other groups. It is undoubtedly paraphyletic, but I have not been able to find an adequate way of splitting it up. Key to species 1. Propodeum with dense golden or silvery to- mentum, completely obscuring sculpture.... 2 - Propodeum at most with sparse golden or silvery tomentum, leaving sculpture easily vis- Iper. ae een el a DI an 3 2. Large species: wing length 26-27 mm; legs largely reddish; New Guinea ....... rex sp. n. — Medium-sized species: wing length 13-19 mm; legs black; Moluccas, New Guinea, Bismarck and Solomon Islands, Australia … HR SURNAM ADS ce ERA EN cognatus Smith 3. Female: outer side of mid tibia only with nor- mal, straight spines apically; male: apical mar- gin of clypeus straight or slightly concave, without median lobe (fig. 17) ................ 4 - Female: outer side of mid tibia with stout, curved spine apically (fig. 64); male: apical margin of clypeus with short median lobe (fig. LS) nent ERA urn 6 4. Wings strongly yellowish to dark brown with violaceous effulgence; India to Moluccas, Phi- lp pinesRraenana PRIA praedator Smith - Wings basically clear, at most slightly yellow- ishiandi'apically infuscate St mr er. 5 5. Pilosity of head and mesosoma dense, yellow- ish brown; face and mesoscutum with dense brownish tomentum obscuring the sculpture; legs reddish; medium-sized species, wing length 19-20 mm; New Guinea ................. “pin ho Belcher ee staudingeri Gribodo - Pilosity of head and mesosoma white to pale 26 yellow; mesoscutum without dense tomentum, sculpture easily visible; legs black; small spe- cies, wing length 10.5-13.5 mm; Sumba, Timor NW AUS tra la ee RE Re rol BM) PRADA formosellus Van der Vecht 6. Wings dark brown with bluish effulgence, legs dark; female: sixth tergite with pygidium (fig. 65); male: 3rd to 7th flagellomere with broad placoids (fig. 45); Burma to Sumatra, Borneo Lei rara ee DI walshae sp. n. - Colour of wings and legs variable; female: sixth tergite without pygidium; male: 3rd flagello- mere rarely with placoid (only in specimens from Philippines and Java) (fig. 44); Sri Lanka to/Philippines and Tlimor tE ana debe Mione subtruncatus Dahlbom Sphex cognatus Smith Sphex cognatus Smith, 1856: 248. Holotype 9: Australia (BMNH!!). Sphex amator Smith, 1856: 246. Holotype @: Australia (OUM!). Sphex formosus Smith, 1856: 254. Holotype 9: Indone- sia, Seram (BMNH!!). Sphex opulentus Smith, 1856: 250. Holotype &: Austra- lia, Richmond River (BMNH!!). Distribution. - Indonesia (Maluku, Irian Jaya), Papua New Guinea, Bismarck Islands, Solomon Islands, Admiralty Islands, Australia. Note. - Specimens from the North Moluccas (Halmaheira etc.) differ from the remaining popu- lations in having silvery tomentum with a faint golden tinge, in particular on the propodeum. Sphex formosellus Van der Vecht Sphex formosellus Van der Vecht, 1957: 366. Holotype Q: Timor (RMNH!!). Distribution. - Indonesia (Timor, Sumba), W. Australia. Sphex praedator Smith (fig. 17) Sphex praedator Smith, 1858: 14. Holotype &: Indonesia, Sulawesi (OUM!). Sphex tyrannicus Smith, 1860: 122. 9: Indonesia, Bacan (type lost, Van der Vecht 1973). Sphex luteipennis Mocsary, 1883: 33. Lectotype 9 (desig- nated by Van der Vecht 1973): Indonesia, Ambon (TMB!) Sphex nigripes var. calopterus Kohl, 1890: 168, 422 [as var. 5]. 6: Indonesia, Sulawesi (NMW). Sphex nigripes var. kohlianus Strand, 1913: 81 [new name for S. nigripes var. calopterus Kohl]. Sphex nigripes var. celebesianus Strand, 1913: 81. 9, &: Indonesia, Sulawesi (NMw) [name for S. nigripes var. 2 of Kohl (1890)]. Sphex nigripes var. melanopodus Strand, 1915: 89. Lec- totype 9 (designated by Van der Vecht 1973): Sri Lanka, Nalanda (NMw!) Distribution. - India and Sri Lanka to China and Malaysia, Indonesia (Sumatra, Sangihe Islands, Java, Sulawesi, Moluccas), Philippines. Sphex rex sp. n. (figs. 43, 61-62) Type material. - Holotype 9, Papua New Guinea, Milne Bay, KBMission, 5.iii.1944, K.V. Krombein (USNM). — Paratype: Indonesia, Irian Jaya, Ifar, 350 m, ii.1957, J. v.d. Assem (1 @, RMNH). Diagnosis This species is immediately recognized by its large size (body length 35 mm), reddish legs and dense golden tomentum on most part of meso- soma. In these respects it is similar only to S. basilicus from Australia, from which it differs as follows: petiole short, 0.35 times as long as hind basitarsus (0.5 in basilicus); 7th sternite of male with proximal transverse depression, 8th sternite trilobate (bilobate in basilicus); anterior femora of female ventrally covered with tomentum (with sparse erect pilosity in baszlicus). Description Female. - Body length 36 mm, length of forewing 26 mm. Black, the following parts reddish: mand- ibles except base and apex, femora except basal half, tibiae, tarsi, tegulae except medially, sixth meta- somal segment. Wings almost clear, apically infus- cate. Pilosity yellow, on vertex and mesoscutal disc infuscate; head and mesosoma with conspicuous dense golden tomentum, completely obscuring sculpture of metanotum and propodeum, tomen- tum on anterior part of mesoscutal disc and scutellar disc dark brown; fore femora ventrally with tomen- tum and pilosity. Clypeus 0.85 times as long as IOC, with blunt median carina on dorsal half, apical margin with short median lobe; 10v 0.85 times IOC; labrum with median carina, which projects apically into a weak tooth; relative lengths of flagellomeres I-III 1 : 0.6 : 0.6. Mesosoma microsculptured, punc- tulate to punctulate-reticulate to shagreened (obs- cured by vestiture almost throughout); scutellum with median longitudinal impression; fore metatar- sus with 8 rake spines, median spines 1.5 times metatarsal width, apical spine as long as second tarsomere; mid tibia with oblique, slightly curved spine apically. Petiole 0.35 times as long as hind basitarsus; sixth tergite without pygidium. HENSEN: Malesian Sphecina 62 63 65 Figs. 62-65. Sphex. 62, S. rex, Q, fore basitarsus; 63-65, S. walshae, 2. 63, fore basitarsus; 64, distal part of mid tibia, outer side; 65, sixth tergite. Male. - Body length 35 mm, length of forewing 27 mm. Like the female except for the following. Mandibles black, mid and hind femora nearly en- tirely black; 7th metasomal segment and 8th ster- nite reddish. First tergite and sides of 6th and 7th with conspicuous yellow pilosity. Clypeus 0.95 times as long as IOC; 10v 0.9 times IOC; labrum ecarinate; relative lengths of flagellomeres I-III 1: 0.65 : 0.65, 5th flagellomere with narrow concave placoid; sixth sternite apically shallowly emarginate, seventh sternite proximally transversely depressed, 8th sternite trilobate, middle lobe curved upward. Etymology Rex is a noun in apposition, from Latin rex = king. Sphex staudingeri Gribodo Sphex staudingeri Gribodo, 1894: 3. Holotype 4: New Guinea (MCG!). Distribution. - New Guinea. 27 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Sphex subtruncatus Dahlbom (figs. 44, 60) Sphex subtruncatus Dahlbom, 1843: 25. Q: ‘Africa’ (Mu- seum Lund!) [probably from NE. India, see Van der Vecht (1973)]. Sphex nigripes Smith, 1856: 254. Syntypes 9: China: Shanghai (BMNH!), Sumatra [non Fabricius, 1793]. Sphex siamensis Taschenberg, 1869: 413. 9: ‘Siam’ (Mus. Univ. Halle!) [probably from the Philippines, accord- ing to Van der Vecht (notes) |. Sphex erythropodus Cameron, 1889: 108. Syntypes 9: India (Mus. Calcutta). Sphex sulciscuta Gribodo, 1894: 2. Holotype 9: Philip- pines, Mindoro (MCG!). Sphex nigripes var. tsingtauensts Strand, 1916: 81A (11): 107. Holotype @: China, Tsingtau [Qingdao] (DEI!!). Sphex subtruncatus coraxus Van der Vecht, 1957: 365. Holotype 9: Indonesia, Sumba, Pogobina (NMB!). Sphex subtruncatus orius Van der Vecht, 1957: 366. Holo- type 6: Indonesia, Flores, Rana Mese (NMB!). Sphex subtruncatus xuthus Van der Vecht, 1957: 364. Holotype g: Indonesia, Sumba, Bondo Kodi (NMB!). Sphex subtruncatus krombeini Van der Vecht in Krom- bein, 1984: 28. Holotype 9: Sri Lanka, Kanneliya Jungle (USNM!). Distribution. - Sri Lanka and India to China and Malaysia, Indonesia (Sumatra, Bangka, Java, Kali- mantan, Lombok, Flores, Sumba, Timor), Philip- pines. Note. - S. subtruncatus exists in a number of geographic forms, which differ in colour (legs black to reddish, wings dark brown to clear), as well as in the number of placoids on the male flagellum. Sphex walshae sp. n. (figs. 18, 45, 63-65) Type material. - Holotype 9, Indonesia, S. Sumatra, 250 m, Bengkulu, Muara Tenam, 16-23.vii.1935, M. E. Walsh (RMNH). — Paratypes: India: Sikkim, Ari to Gna- tong, 7-2500 ft, iv.1894, Bingham (1 9, USNM). Burma: Tenasserim, Ihaungyin Valley, v.1891, Bingham (1 9, BMNH). Malaysia: Malay peninsula, Kedah Peak, 3000- 3500 ft, 15.111.1928, H. M. Pendlebury (1 9, BMNH); Sa- rawak, C. J. Brooks (1 9, BMNH); Sabah, Sandakan, Baker (2 9, USNM); Sabah, 19 km N. of Kalabakan, 19.xi.1962, Y. Hirashima (1 ©, BPBM); North Borneo, Frühstorfer (1 Q, TMB). Indonesia. Sumatra: Anei kloof, 500 m, E. Jac- obson (3 9, RMNH); Bengkulu, 50 m, 10-18.v.1935, M. E. Walsh (1 9, RMNH); Bengkulu, Bukit Item, 650 m, 11- 15.vi.1935, M. E. Walsh (1 9, RMNH); Bengkulu, Tandjong Sakti, 1-10.vi.1935, M. E. Walsh (2 & 1 9, RMNH); Lubuk Sikaping, 450 m, L. Lundeshagen (1 9, RMNH), Aur Ke- manis, E. Jacobson (1 9, RMNH); no locality (1 9, Zoolo- gische Staatssammlung, Miinchen). Kalimantan: Keta- pan, v.1937, M. E. Walsh (2 9, RMNH); Sampagan, 0 m, 1927, M. E. Walsh (1 4, RMNH); South coast (1 9, RMNH); Kembang Djangut, 75 m, 24-30.xi.1956, A.M.R. Wegner (5 9, RMNH); no locality, Muller (1 & 5 9, RMNH). ‘Ind. or.?’ (1 9, RMNH). 28 Diagnosis This species is similar to other large, dark- winged, dark-legged forms: the representative of S. subtruncatus in the Philippines and the represen- tative of S. praedator in the Moluccas (apart from species belonging in other groups). As these forms are allopatric, S. walshae may be separated from sympatric forms on the basis of the dark brown wings with violaceous effulgence. Unlike S. prae- dator, S. walshae has a stout curved spine at the outer end of the female mid tibia, and the male clypeus has a median apical lobe. In these respects S. walshae is similar to S. subtruncatus, from which it differs in having a pygidial plate in the female, and placoids on third to 7th flagellomere in the male (variable in S. subtruncatus, the mentioned condition occurring in specimens from the Philip- pines and Java). Description Female. - Body length 23-28 mm, length of fore- wing 17-20 mm. Black. Wings dark brown with violaceous effulgence. Pilosity of head and meso- soma black; clypeus and frons with sparse silvery tomentum, mesosoma with sparse, inconspicuous dark brown tomentum; fore femora ventrally with sparse erect hairs, no tomentum. Clypeus 0.8 times as long as IOC, without median carina, apical mar- gin slightly raised and with a pair of short teeth medially; rov 0.8 times 10C; labrum with median carina, which projects apically into a tooth; relative lengths of flagellomeres I-III 1 : 0.7 : 0.75. Meso- soma microsculptured: punctulate-reticulate to shagreened, mesonotum and mesopleuron largely shallowly punctate, punctures 1-2 diameters apart; propodeum finely rugose; scutellum with median longitudinal impression; fore basitarsus with 9-10 rake spines, median spines 1.5 times tarsal width, apical spine as long as second tarsomere; apex of mid tibia with stout curved spine. Petiole 0.4 times as long as hind metatarsus; sixth tergite with pyg- idium. Male. - Body length 21-24 mm; length of fore- wing 16-18 mm. Like the female except for the following. Clypeus 0.9 times as long as IOC, apical margin with short, depressed median lobe; 10v 0.85 times 10C; labrum ecarinate; length of flagello- meres I-III 1: 0.75 : 0.75, third to seventh flagello- mere with broad, flat placoid. Seventh sternite weakly emarginate; eighth sternite triangular. Etymology Walshae is a noun in genitive case, refering to Mrs. M. E. Walsh, collector of a substantial part of the type series. REFERENCES Berland, L., 1928. Les Sphegidae du Muséum National de Paris. - Bulletin du Muséum d'Histoire Naturelle de Paris 34: 331-333. Bingham, C. T., 1890. On new and little known Hyme- noptera from India, Burma, and Ceylon. - Journal of the Bombay Natural History Society 5: 233-252. Bingham, C. T., 1894. On new and little known Hyme- noptera from India, Burma, and Ceylon. - Journal of the Bombay Natural History Society 8: 358-390, 2 pls. Bohart, R. M., & A. S. Menke, 1976. Sphecid wasps of the world, a generic revision: i-ix, 1-695. - University of California Press, Berkeley. Cameron, P., 1889. Hyme- noptera Orientalia I. - Memoirs and Proceedings of the Manchester Literary and Philosophical Society (4) 2: 91-152. Cameron, P., 1901. On the Hymenoptera collected in New Britain by Dr. Arthur Wiley. - Proceedings of _ the Zoological Society of London 1901: 224-248. Cameron, P., 1902. Descriptions of new genera and spe- cies od Hymenoptera from the Oriental zoological region. - Annals and Magazine of Natural History (7) 9: 145-155, 204-215, 245-255. Cameron, P., 1906. A fourth contribution to the know- ledge of the Hymenoptera of Sarawak. - Journal of the Straits Branch of the Royal Asiatic Society 46: 93-168. Christ, J. L., 1791. Naturgeschichte, Classification und Nomenclatur der Insecten vom Bienen- Wespen- und Ameisengeschlechte: 1-535, pls. 1-60. - Frankfurt. Costa, A., 1864. Elenco delle specie di animali immesse per aquisti. - Annuario del Museo Zoologico della R. Universita di Napoli 2 (1862): 111-112. Dahlbom, A. G., 1843. Hymenoptera europaea, praecipue borealia, etc., 1 (1): 1-172. - Lund. Dalla Torre, C. G. de, 1897. Catalogus Hymenopterorum hucusque descriptorum sytematicus et synonymicus. 8, Fossores: i-viii + 1-749. - G. Engelmann, Lipsiae. Fabricius, J. C., 1787. Mantissa Insectorum etc., 1: 1-20 + 1-348. - Hafniae. Fabricius, J. C., 1793. Entomologia systematica, 2: 1-8 + 1-519. - Hafniae. Fabricius, J. C., 1804. Systema piezatorum: 1-14 + 1-440 + 1-30. - Brunsvigae. Gribodo, G., 1894. Hymenopterorum novorum diagnoses praecursoriae. - Miscellanea Entomologica 2: 2-3. Guérin-Méneville, F. E., 1831, 1838. Crustacées, Arach- nides et Insectes. In: L. F. Duperrey, Voyage autour du monde sur la Coquille. II (2) (1838), pls. (1831). Guillou, E. J. F. le, 1841. Catalogue raisonné des insectes Hymenopteres recueillis dans le voyage de circumnav- igation des corvettes l’Astrolabe et la Zelée. - Annales de la Société Entomologique de France 10: 319-320. Kohl, F. EF, 1885. Die Gattungen der Sphecinen und die paläarktischen Sphex-arten. - Természetrajzi Fiizetek 9: 154-207. Kohl, F. F, 1890. Die Hymenopterengruppe der Sphe- cinen. I: Monographie der natiirlichen Gattung Sphex Linné (sens. lat.). - Annalen des Naturhistorischen Museums in Wien 5: 77-104, 317-461. Kohl, F. F,, 1895. Zur Monographie der natürlichen Gat- tung Sphex Linné. - Annalen des Naturhistorischen Museums in Wien 10: 42-774, pls. IV, V. Kohl, F. F., 1898. Ueber neue Hymenopteren. - Termés- zetrajzi Füzetetek 21: 325-367, pl. XV. Krombein, K. V., 1984. Biosystematic studies of Ceylo- HENSEN: Malesian Sphecina nese wasps, XII: Behavioral and life history notes on some Sphecidae. - Smithsonian Contributions to Zoology 387: 1-30. Lepeletier de Saint-Fargeau, A., 1845. Histoire naturelle des insectes Hyménoptères, 3: 1-644. - Roret, Paris. Matsumura, S., 1912. A thousand insects of Japan, suppl. 4: 1-247, pls XLII-LV. Mocsary, A., 1883. Hymenoptera nova europaea et exo- tica. - Ertekezézek a Természettudomanyok Köreböl 572: Rohwer, S. A., 1921. The Philippine wasps of the subfam- ily Pseninae. - Philippine Journal of Science 19: 665- 676. Saussure, H. de, 1867. Hymenoptera. In: Reise der öster- reichische Frigatte Novara, etc., Zoologischer Theil, 2 (1): 1-156, pls I-IV. - Wien. Saussure, H. de, 1869. Hyménoptères divers du Musée Godeffroy. — Stettiner Entomologische Zeitung 30: 53-64. Smith, F., 1856. Catalogue of the Hymenopterous insects in the British Museum. Sphecidae, Larridae and Cra- bronidae: 207-497. - London. Smith, F., 1858. Catalogue of the Hymenopterous insects collected at Sarawak, Mount Ophir and at Singapore by A. R. Wallace. - Journal of Proceedings of the Linnean Society of London, Zoology 2: 43-130, 2 pls. Smith, F., 1858. Catalogue of the Hymenopterous Insects collected at Celebes by Mr. A. R. Wallace. - Journal of Proceedings of the Linnean Society of London, Zool- ogy 3 (1859): 4-27. Smith, F., 1859. Catalogue of the Hymenopterous insects collected by Mr. A. R. Wallace at the islands of Aru and Key. - Journal of Proceedings of the Linnean Society of London, Zoology 3: 132-178. Smith, F, 1860. Descriptions of Hymenopterous insects collected by Mr. A. R. Wallace in the islands of Bat- chian, Kaisaa, Amboyna, Gilolo, and at Dory in New Guinea. - Journal of Proceedings of the Linnean So- ciety of London, Zoology 5: 93-143. Smith, F, 1862. Catalogue of Hymenopterous insects col- lected by Mr. A. R. Wallace in the islands of Ceram, Celebes, Ternate and Gilolo. - Journal of Proceedings of the Linnean Society of London, Zoology 6: 36-66, 1 pl. Smith, F, 1873. Hymenoptera. In: J. L. Brenchley, Jottings during the cruise of H. M. S. Curagao, etc. - London. Strand, E., 1910. Drei neue Crabroniden nebst Bemerkun- gen zur Verbreitung einiger anderen Hymenopteren, etc. - Jahrbücher des Nassauischen Vereins für Natur- kunde 63: 46-52. Strand, E., 1911. Crabronidae und Pompilidae. - Abhand- lungen der Senckenbergischen naturforschenden Ge- sellschaft 34: 223-234. Strand, E., 1913. H. Sauter’s Formosa Ausbeute. Crabro- nidae und Scoliidae I. - Archiv für Naturgeschichte 79A (3): 76-87. Strand, E., 1915. Ueber einige orientalische und paläark- tische Crabroniden der Gattungen Sphex, Sceliphron und Ammophila im Deutschen Entomologischen Mu- seum. — Archiv für Naturgeschichte 81A (5): 88-97. Strand, E., 1916. Ueber einige exotische Crabroniden der Gattungen Sphex und Sceliphron im Deutschen En- tomologischen Museum. - Archiv für Naturge- schichte 81A (11): 98-109. Taschenberg, E., 1869. Die Sphegidae des zoologischen Museums der Universität in Halle. - Zeitschrift für 29 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 die gesammte Naturwissenschaft 34: 407-435. Turner, R. E., 1908. Note on the Australian wasps of the family Sphegidae, with descriptions of new species. — Proceedings of the Zoological Society of London 1908: 457-535. Turner, R. E., 1910a. Additions to our knowledge of the fossorial wasps of Australia. — Proceedings of the Zoological Society of London 1910: 252-355. Turner, R. E., 1910b. New fossorial Hymenoptera from Australia. - Transactions of the Entomological Society of London 1910: 407-429, pl. I. Turner, R. E., 1912. Notes on fossorial Hymenoptera VI. On the species collected in New Guinea by the expe- dition of the British Ornithologists Union. - Annals and Magazine of Natural History (8) 9: 194-202. Turton, W., 1800. System of nature through the three grand kingdoms, etc. I: 1-784. - D. Williams, Swansea. Vachal, J., 1908. Deuxiéme note sur les Hyménopteres de la Nouvelle Calédonie. - Revue d’entomologie, Caen 26: 113-123. 30 Vecht, J. van der, 1957. The Sphecoidea of the Lesser Sunda Islands (Hym.). I. Sphecinae. - Verhandlungen der Naturforschenden Gesellschaft in Basel 68: 358- 372. Vecht, J. van der, 1961. Hymenoptera Sphecoidea Fabri- ciana. - Zoologische Verhandelingen 48: 1-85. Vecht, J. van der, 1973. Contribution to the taxonomy of the Oriental and Australian Sphecini (Hymenoptera, Sphecoidea). - Proceedings Koninklijke Nederlandse Akademie van Wetenschappen (C) 76: 341-353. Vecht, J. van der, & K. V. Krombein, 1955. The subspecies of Sphex sericeus (Fabr.) (= S. aurulentus auct., nec Fabr., 1787) (Hymenoptera, Sphecidae). - Idea 3: 33- 43. Received: 3 November 1990 Accepted: 7 January 1991 P. HUEMER Tiroler Landesmuseum Ferdinandeum Innsbruck CARYOCOLUM DELPHINATELLA (CONSTANT) SP. REV, A SENIOR SYNONYM OF C. FIORII (KLIMESCH) (LEPIDOPTERA: GELECHIIDAE) Huemer, P., 1991. Caryocolum delphinatella (Constant) sp. rev., a senior synonym of C. fioru (Klimesch) (Lepidoptera: Gelechiidae). - Tijdschrift voor Entomologie 134: 31-34, figs. 1-7. [ISSN 0040-7496]. Published 1 July 1991. Caryocolum delphinatella (Constant) is recognized as good species and revocated from synonymy with C. interalbicella (Herrich-Schäffer). The female genitalia are described and figured and data on biology are given. C. fioriti (Klimesch) is synonymised with C. delphinatella. P. Huemer, Tiroler Landesmuseum Ferdinandeum, Museumstr. 15, A-6020 Innsbruck, Austria. Key words. - Gelechiidae, Caryocolum, synonymy. In 1890 the well-known French lepidopterist A. Constant described Lita delphinatella from a single female collected in July in the Alpes du Dauphiné. Klimesch (1954) in his revision of European spe- cies of the former genus Lita feeding on Caryophyl- laceae, queried delphinatella as a synonym of Ca- ryocolum interalbicella (Herrich-Schäffer). This synonymisation was accepted by the present author (Huemer 1988) although de/phinatella was treated as a separate species in the French list (Leraut 1980). C. frorız was described from five males col- lected in the Abruzzi (Klimesch 1953). Later the species was also recorded from the Swiss and French Alps. When re-valuating the original description of delphinatella, including the colour plate depicting the holotype, it became clear that this species is not a synonym of :interalbicella but conspecific with fioru. This assumption is further supported by rich material, including two females, collected in the south-western Alps in July 1990. Caryocolum delphinatella (Constant) sp. rev. Lita delphinatella Constant, 1890a (May 28): 6, pl. 1, fig. 2 [moth in colour]. Holotype 9, France: Alpes du Dauphiné, vii [not traced]. Lita delphinatella Constant, Meess 1910: 370. Phthorimaea delphinatella (Constant); Meyrick 1925: 95; Gaede 1937: 254; Lhomme [1946]: 632. Caryocolum delphinatella (Constant); Leraut 1980: 79. Gnorimoschema fiorii Klimesch, 1953: 277, figs 23, 24. Lectotype 8 (Designated by Huemer 1988: 481), Italy: Abruzzen, Gran Sasso, 25.vii.1935 (Fiori) (coll. Kli- mesch, Linz) [examined]. Syn. n. Caryocolum fiorii (Klimesch); Huemer 1988: 480. Description Adult (Figs 1, 3-5). - Fore wing length & 7.0-7.5 mm, ® 6.5-7.0 mm. Head white; Labial palpus white, third segment mottled with dark brown. Thorax white, tegulae dark brown basally. Fore wing dark brown, mottled with light brown; white transverse fasciae at one-fifth and one-half, me- dially narrowed and often reduced towards costa; fasciae fused and mottled with light brown dor- sally; separate white subcostal spot at about two- fifth always well developed; white costal and tornal spots at four-quarters variable in extension, separ- ate or fused to fascia; fringes whitish with dark brown basal line. Hind wing light grey. Male genitalia (Huemer 1988: Fig. 122). Female genitalia (Figs 6, 7). - Apophyses poste- riores 1.60 mm, apophyses anteriores 0.43 mm; eighth segment without processes; ostium bursae surrounded by folds; antrum short, funnel-shaped, posteriorly fused with pair of short lateral sclero- tizations of ductus bursae which are level with the tip of the apophyses anteriores; corpus bursae py- riform with long, hook-shaped signum. Biology The immature stages are still unknown. Accord- ing to personal observations in France the larvae probably feed on Minuartia sp. and Silene sp. (Ca- 21 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 5 Figs. 1-5. Caryocolum spp. - 1, C. delphinatella, original figure by Constant; 2, C. interalbicella; 3, C. delphinatella (fiorti lectotype); 4, C. delphinatella 9: 5, C. delphinatella 8. 32 ryophyllaceae) around which the adults have been collected in numbers in July and early August; most specimens were attracted to artificial light sources. The females examined were both collected during day time resting on rocks. C. delphinatella is an alpine species which occurs from about 1600 to 2500 m. Distribution Only known from the Abruzzi and the south- western Alps (Alpes Maritimes, Alpes de Haute Provence, Hautes Alpes, Isère (Dauphiné), Wal- liser Alpen, Alpi Cozie); according to Lhomme [1946] also in the Pyrenées (Haute-Garonne). Remarks Despite the efforts of several lepidopterists the holotype of delphinatella could not be traced in MNHN Paris or any other institution; however, Constant’s original description and colour illustra- tion, the latter reproduced here in monochrome (Fig. 1), leave no doubt about the identity of this species. It is well characterized by the pure white head and thorax as emphasised by Constant, and the white subcostal forewing spot at two-fifths (Figs 1,3-5). The synonymisation with Caryocolum interalbicella is certainly incorrect as this species has a whitish to cream head and tegulae and a dark brown thorax (Fig. 2). The few French Caryoco- lum-species with white colour of head and thorax differ from delphinatella: C. saginella (Zeller) is distinctly smaller (4.0-5.0 mm); C. cauligenella (Schmid) has a creamy rather than pure white head and thorax and also broader fore wings without the costal spot present in delphinatella; French spe- cimens of C. peregrinella (Herrich-Schäffer) are characterized by the larger size (7.5-9.0 mm), broader fore wings and extended white markings of the fore wing without separate subcostal spot. C. leucothoracellum (Klimesch) has different fore- wing markings with white subcostal streak and a medial spot. C. frori perfectly matches delphina- tella and is here placed as a junior subjective syn- onym. C. fiorii was recently recorded as new to France (Huemer 1989) because the true identity of delphinatella, at that time still treated as synonym of interalbicella, was not recognized. The female genitalia of delphinatella are similar to those of other species of the trauniella-group. The antrum is distinctly longer than in peregri- nella (Herrich-Schäffer) and trauniella (Zeller). The lateral sclerotizations of the ductus bursae are level with the tip of the apophyses anteriores in delphinatella whereas in other species of the group they are at most half that length. The differences in the male genitalia have been discussed pre- viously (Huemer 1988). HUEMER: Caryocolum delphinatella 4 Figs. 6-7. Caryocolum delphinatella, female genitalia. - 6, eighth segment; 7, signum (X 2 magnification of eighth segment). Material examined (additional to that previously re- corded (Huemer 1988, 1989). — Italy: 4 8, Abruzzen, Prov. Rieti, Terminillo-Paß, 18-2000 m, 5-6.viii.1970, leg. Groß; 2 4, P. d. Fargno, ob. Bolognola, Mti. Sibillini, 16-1700 m, 14.viii.1970, leg. Groß (Löbbecke Museum, Düsseldorf); 1 g, Prov. Torino, Alpi Cozie, Colle delle Finestre, 2150 m, 27.vii.1990, leg. Huemer & Tarmann. France: 39 4, 1 9, Dep. Alpes-Maritimes, Marguareis W- Hang, Navela, 2100-2200 m, 21-23.vii.1990, leg. Huemer & Tarmann; 22 g, 1 9, Dep. Alpes-de-Haute-Provence, SW Castel de Restfond, Roche Chevaliére, 2480 m, 25.vii.1990, leg. Huemer & Tarmann; 10 4, Dep. Alpes- de-Haute-Provence, SW Castel de Restfond, Set. de Caire Brun N-Flanke, 2420 m, 25.vii.1990, leg. Huemer & Tar- mann (Tiroler Landesmuseum Ferdinandeum, Inns- bruck). ACKNOWLEDGEMENTS I gratefully acknowledge the help received for this study from Mr O. Karsholt (ZMUC, Copenha- gen), Dr S. Lòser (Diisseldorf), Dr G. Luquet (MNHN, Paris) and Dr K. Sattler (BMNH, London). Furthermore I am indebted to my colleagues M. Kahlen, B. Plössl and Dr G. Tarmann (Innsbruck) for their help in the field and valuable comments. 33 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 REFERENCES Constant, A., 1890a. Séance du 26 Juin 1889. - Bulletin de la Societe Entomologique de France 1889: 125. Constant, A., 1890b. Descriptions de Microlepidopteres nouveaux ou peu connus. - Annales de la Societe Entomologique de France 69: 5-16. Gaede, M., 1937. Gelechiidae. - Lepidopterorum Catalo- gus 79: 1-630. Huemer, P, 1988. A taxonomic revision of Caryocolum (Lepidoptera: Gelechiidae). - Bulletin of the British Museum (Natural History) (Entomology) 57: 439- Il: Huemer, P., 1989. Bemerkenswerte Funde von Caryoco- lum-Arten aus den Südalpen und dem Mediterraneum (Lepidoptera, Gelechiidae). - Nachrichtenblatt der bayerischen Entomologen 38: 37-40. Klimesch, J., 1953-1954. Die an Caryophyllaceen le- benden europäischen Gnorimoschema Busck (=Phthtorimaea Meyr.)-Arten. — Zeitschrift der Wiener Entomologischen Gesellschaft 38: 225-239, 275-282, 311-319, 39: 273-288, 335-341, 357-362. 34 Leraut, P., 1980. Liste systématique et synonymique des Lepidopteres de France, Belgique et Corse. - Supple- ment à Alexanor et au Bulletin de la Société Entomo- logique de France, 334 pp. Lhomme, L., [1946-1948]. Catalogue des Lépidoptères de France et de Belgique 2: 489-648. - Douelle (Lot). Meess, A., 1910. xxxviii. Familie. Gelechiidae. - In: Hof- mann, E. & A. Spuler (eds). Die Schmetterlinge Eu- ropas. 2, 523 pp. - Stuttgart. Meyrick, E., 1925. Lepidoptera Heterocera. Fam. Gelechi- adae. - Genera Insectorum 184: 1-290, pls 1-5. Received: 23 November 1990 Accepted: 26 November 1990 I. LANSBURY Oxford University Museum CUTICULAR BLADES AND OTHER STRUCTURES OF DIAPREPOCORIS KIRKALDY AND STENOCORIXA HORVATH (HETEROPTERA: CORIXIDAE) Lansbury, I., 1991. Cuticular blades and other structures of Diaprepocoris Kirkaldy and Stenocorixa Horvath (Heteroptera: Corixidae). - Tijdschrift voor Entomologie 134: 35- 46, figs. 1-27. [ISSN 0040-7496]. Published 1 July 1991. Diaprepocoris Kirkaldy and Stenocorixa Horvath possess cuticular blades and modified swimming hairs on their hind tibiae and tarsi which resemble those of gyrinids (Coleo- ptera: Gyrinidae). These have not previously been recorded in the Corixidae. Unusual macrotrichia are also present on the clavus and corium of all known species of both genera. The unique characters of these two genera support their separate subfamily placements. I. Lansbury, Hope Entomological Collections, University Museum, Oxford, OX1 3 PW, UK. Keywords. - Australia, Africa, Cuticular blades, modified swimming hairs, respiration, Corixidae. A study of the Australasian Diaprepocoris has revealed the presence of cuticular blades and modi- fied swimming hairs on the hind tibiae and tarsi. These structures have not previously been reported in the Corixidae. A survey of the corixid genera has shown that a similar combination of cuticular blades and modified swimming hairs are also pres- ent in the genus Stenocorixa which is restricted to tropical Africa. The cuticular blades and associated specialised swimming hairs are described and their probable function discussed in relation to the nek- tonic (submerged) swimming strategies and respi- ration in both subfamilies. The double hair pile layer on the forewings of Diaprepocoris is figured and a dual function sug- gested for the hair pile. Diaprepocoris is compared with Stenocorixa, the later apparently having smooth forewings. The relationship between the gas stores of Diaprepocoris and the specialised swimming hairs of the hind legs are discussed. No data are at present available for Stenocorixa. The taxonomic position (isolation) of the two subfam- ilies is briefly discussed. Diaprepocorinae Lundblad, 1928 Single extant genus Diaprepocoris Kirkaldy, 1897, type species D. barycephala Kirkaldy, 1897. Within the Diaprepocorinae, Popov (1971) in- cludes Gazimuria scutellata Popov, from the lower Jurassic, Akutaev Series, Chitin Province, East Transbaikals and Karataviella brachyptera Becker- Migdisova from upper Jurassic, Chayan Region, Chimkent Province, South Kazakhstan. Walton (1940) split the Diaprepocorinae into two tribes using the structure of the female pala: Diaprepocorini. - Female pala digitform, few sieve hairs (about 10) and short. Type/Genus spe- cies Diaprepocoris barycephala. Corixanectini. - Female pala short and broad, inner side flattened and fringed with numerous long sieve hairs. Type/Genus species Corixanecta zealandiae Hale. Stenocorixinae Hungerford, 1948 Single extant genus, Stenocorixa Horvath, 1927, type species S. protrusa Horvath, 1927, monobasic. Jaczewski (1928) placed Stenocorixa in the Corixinae; Poisson & Jaczewski (1928) discussing the systematic position of Stenocorixa concluded that the poorly developed abdominal asymmetry, shape of the pala claw and other features suggested a relationship with Cymatia Flor. They ended by stating ‘The generic distinctness of Stenocorixa Horv. is beyond any doubt and even as a separate genus it seems to stand rather apart of the other genera of the subfamily Corixinae’. 35 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 SN: Figs. 1-4. Diaprepocoris barycephala Kirkaldy. - 1, corium, double hair pile, note collapsed microtrichia and longi- tudinal ridges on macrotrichia X3200; 2, inner lateral margin of clavus, note ‘shephards crook' macrotrichia X400; 3, middle of corium X800; 4, membrane of elytra, note more erect microtrichia and enlarged socket of macrotrichia X6400. Reduction 84%. 36 DESCRIPTIVE PART Diaprepocoris, structure trichia Macrotrichia of the clavus and corium varying in shape. The hairs along the inner margin of the corium are apically curled over forming a ‘shep- herds crook’ (fig. 2) and are longer and thinner than those on the inner areas adjacent to the clavus (fig. 3). The microtrichia are densely packed and resemble a plastron-like layer however, as they have collapsed and become matted, they clearly do not function as a plastron. The distribution of the microtrichia is fairly uniform over the clavus and corium. The membrane differs in that the micro- trichia do not seem to have collapsed as much as those on the clavus and corium (fig. 1 clavus; fig. 4 membrane). The shape of the sockets within the microtrichia from which the macrotrichia arise,’ structurally suggests that the latter can be elevated and depressed (fig. 1). The macrotrichia are rather more slender and fewer in number on the mem- brane. Those on the elytra are longitudinally ridged (fig. 1). The division between the double hair layer and the edge of the forewing is distinct (fig. 5). The lateral margin at the nodal furrow shows a scale- like appearance with no hair layers (fig. 6). Stenocorixa, structure trichia Unlike nearly all other genera of corixids, the pronotum and hemelytra are not rastrate, but smooth and shining with scattered long strap-like hairs arising from angled depressions, each with a complete basal ring (fig. 13). Caudad of each de- pression, an elongate channel within the clavus- corium, the strap-like hairs are almost certainly lodged in the channel(s) when the corixed is swim- ming. Diaprepocoris, cuticular blades and swimming hairs The hind tarsi of Diaprepocoris have in addition to the rows of swimming hairs, fringes of cuticular blades as long or longer than their respective tarsal segments. When not in use, they overlie each other like a venetian blind (fig. 7). Fanned out they in- crease the surface area of the tibiae-tarsi. The spread of the blades increases the surface area of the Ist tarsus along the leading edge by 3.5 X 10 (this and following figures approximate) - trailing edge has a greater density of fine swimming hairs overlying a row of blades increasing the surface area by 10 X 2. The leading edge of 2nd tarsus has a row of blades decreasing in length distally, in- creasing surface area by 3.0 x 10 — trailing edge has LANSBURY: Cuticular blades in Corixidae fewer swimming hairs, the blades are roughly all the same length, increasing surfaces area by 2.0 10. The cuticular blades thus greatly increase the surface area (figs. 17, 18). According to Nachtigall (1974) cuticular blades create about 90% thrust of that of a solid object of equal size in the Gyrinidae (Coleoptera). The blades of Diaprepocoris appear to have a function similar to those of gyrinids, the latter also use their middle legs when swimming. The hind coxae of Diaprepocoris conform to the usual co- rixid form. The ventral surface of the coxae is flattened to accommodate the trochanter-femur in the full recovery position, the femur forming a 33° angle with the midline of the body. The flattened area is partially enclosed by the tip of the meta- xyphus and prevents excessive rotation of the hind leg (fig. 19). Schenke (1965, 1966) gives an account of a corixid swimming using Corixa punctata Illiger as a model. There are striking differences between the two genera. Corixa has a complex arrangement of rows of hairs, each with an incomplete basal ring, this enables the hairs to be folded flat against the leg during the recovery stroke. The tarsal hairs are supported by a row or rows of stout flexible spines which function in a similar fashion. There are few hind tarsal spines on Diaprepocoris com- pared with Corixa; D. barycephala \st tarsus has between 10-13 spines in each of three lateral rows, one either side of the trailing edge and one along the leading edge. All the spines are semi-erect and probably rotate in much the same way as Corixa. The fringes of swimming hairs are arranged in a distinct manner. According to Schenke (1965) there are in excess of 5000 hairs on each rowing leg of Corixa. In Diaprepocoris, the leading edge of the lst tarsus has about 30 long hairs, each having an incomplete basal ring (fig. 8) and a larger number along the trailing edge. Second tarsus with 8 long hairs along the leading edge and about 30 along the trailing edge. Figure 9 shows the incomplete basal rings of the cuticular blades in the recovery posi- tion, fig. 10 shows the folding of the blades on the 2nd tarsus partially overlapping the apical claws and extreme length of some of the blades. The hind tibiae moderately spinose (fig. 23) interspersed with hairs, some are expanded distally with a series of fine incisions (fig. 11). These modified hairs may reduce drag and loading on the hind legs, another function may be to move expired gas backwards over the elytra prior to the corixid surfacing to renew its gas store. The distribution of the explan- ate hind tibial hairs and cuticular blades of the tarsi are not uniform throughout the genus, D. baryce- phala (fig. 17), D. personata Hale (fig. 18) and D. pedderensis Knowles (figs. 20, 21). There is also variation in the development of the swimming SY TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 | | Figs. 5-8. Diaprepocoris barycephala Kirkaldy. - 5, outer lateral margins of corium and edge of double hair pile layer X3200; 6, lateral margin of forewing at nodal furrow with scale-like surface X800; 7, overlapping cuticular blades on hind tarsus X200; 8, hind tarsi, note incomplete basal rings X400. Reduction 84%. 38 LANSBURY: Cuticular blades in Corixidae Figs. 9-12. Diaprepocoris barycephala Kirkaldy. - 9, base of cuticular blades showing the incomplete basal rings X800; 10, cuticular blades on hind tarsi, note extreme length of blades X50; 11, incised hairs on the hind tibia X800; 12, immature Diaprepocoris, hind tarsus X50. Reduction 84%. 39) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 14, ind tarsi, note length and density of blades X50; 16, ’ ith single strap-like hair, note channel in elytra X3200 Figs. 13-16. Stenocorixa protrusa Horvath. - 13, elytra w explanate hind tarsi with cuticular blades X24; 15, explanate h hind tarsi X400. Reduction 84%. d hairs on INCISE 40 hairs on the middle legs; in pedderensis none are visible on a slide mounted leg (fig. 22), personata has about 20 hairs visible on the tibia and barycephala has about 50 hairs on the tibia and two rows of about 60 hairs on the tarsus. Stenocorixa, cuticular blades and swimming hairs The hind tarsi of Stenocorixa (fig. 14) are the most explanate of all the corixids, they most closely resemble those of Lethocerus (Belostomatidae, He- teroptera). The cuticular blades are extremely long forming a dense matted fringe (fig. 15). It has not been possible to determine from available prepa- rations if each of the blades have an incomplete basal ring q.v. Diaprepocoris (fig. 9). The long swimming hairs fringing the rows of blades each appear to have incomplete basal rings. The trailing edge of the Ist tarsus has about 10 groups of stout semi-erect spines, between each group, a single thicker spine. The rows partially enclose several rows of long fan-shaped hairs which are deeply incised distally (fig. 16), it is likely that these hairs have a similar function to those of Diaprepocorts. The fine structure of the clavus and corium of Stenocorixa does not suggest that it has a large supra-alar gas store, its overall streamlined appear- ance is totally different from that of Diaprepocoris. The complete basal rings would prevent the hairs laying flat against the forewings and would pre- sumably cause minor eddies over the dorsum of the swimming corixid. The hairs are too few in number to act as a hydrofuge layer and may have some sensory function or be used for manouvering nek- tonically. There is a rather greater degree of flexibility between the hind tibiae-tarsi than in Dia- prepocoris. The hind tibiae of Stenocorixa have a pair of prominent projections distally. The acu- minate insertion of the tarsus within the tibia sug- gests that there is considerable flexibility which would increase the corixids manouverability. The 2nd tarsus is capable of partial folding as in the Gyrinidae, this would be advantageous in the re- covery stroke reducing drag. The hind legs of Stenocorixa are pro-rata much longer than those of Diaprepocoris. Diaprepocoris: hind tibiae not explanate and not reaching the end of the abdomen. Stenocorixa: hind femur-tibia-tarsus explanate, tibia reaching the end of the abdomen. The power stroke of Stenocorixa may exert a greater propulsive effort, the folding hind tarsi reducing drag during the recovery stroke. The rel- ative shortness of the hind legs of Diaprepocoris with possible reduction of energy in the power LANSBURY: Cuticular blades in Corixidae stroke may be compensated for by the supra-alar gas store, this would enable the corixid to stay submerged for longer periods, thus avoiding pre- dators. The extra bouyancy would however, also mean that the corixid would need to keep up a high level of power strokes to maintain depth and atti- tude when submerged and not resting on sub- merged objects. DISCUSSION Thorpe & Crisp (1949) group aquatic animals with hydrofuge hairs in four categories, placing Corixa (Corixinae) in group III, members of which must surface at intervals to renew their gas stores. They are bouyant rather than heavier than water and often have a double hair pile. Diaprepocoris according to Parsons (1976) has a much larger exposed dorsal air store covering the scutellum and forewing; the Micronectinae Jac- zewski have the supra-alar store restricted to the anterior part of the embolium and the anterior- most part of the claval suture; the Corixinae Jac- zewski have a slightly larger air store extending the length of the embolium, anterior part of the clavus suture and that part of the clavus covered by the pronotum. The mesonotal air store of the Micro- nectinae and Corixinae covered by the pronotum. As a form of air bubble respiration, Diaprepocoris has a much larger gas store than most other corix- ids, in addition to the supra-alar store, it has sub- alar gas stores beneath the forewings and over much of the venter. The presence of large supra-alar air stores and sub-alar stores may increase the bouyancy of Diaprepocoris, the bouyancy levels possibly being related to the density of swimming hairs and cutic- ular blades on the hind legs. Possible secondary functions of the external gas layer may be either as a flexible skin which would enable the corixid to swim with less friction. With the exception of D. pedderensis, all Diaprepocoris species are rela- tively short and squat, their greatest width being just over half total length. An alternative function may be acoustic; Theiss (1982) has shown that in Corixa, the cervical air store i.e. air stores formed by the posterior and lateral parts of the head being flattened forming flanges which overlap the pro- thorax, are used in the generation and radiation of sound, thus the dual function of air stores has been demonstrated. Taxonomy Diaprepocoris and Stenocorixa are the most dis- tinctive genera of the extant Corixidae. Characters common to both subfamilies are: 41 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 WN . — 4 Figs. 17-18. Diaprepocoris hind tarsi. - 17, D. barycephala Kirkaldy, scale line 1 mm, 18, D. personata Hale, scale line 0.25 mm. - Hind tarsi fringed with cuticular blades - Little difference between the male and female palae The Diaprepocorinae share a set of characters which distinguishes the subfamily from all others: 1. Ocelli on vertex between compounds eyes (fig. 24) Fore tibia-tarsus fused (fig. 25) 3. Tarsal claws of hind legs apical (figs. 17, 18 and 21) Muscle attachment to male genital capsule dorsal Female ovipositor with a recognisable blade- like structure The Stenocorixinae share a limited set of char- acters which distinguish the subfamily: 1. Clavus and corium smooth without microtri- chia (fig. 13) 2. Clavus and corium with scattered long strap- like hairs 3. Abdominal asymmetry of male 6-7th tergites hardly differing from female 4. Male left paramere absent Various proposals have been made regarding the validity of the corixid subfamilies. Leston (1955) partially summarised the proposals of China (1943) and Hungerford (1948) and observed that most authors treat the Corixidae as a monophyletic group. Popov (1971) altered this concept to include the Shurabellidae (Shurabella lepyronopsis Becker- Migdisova, Liassic fossil). The Corixoidea should be considered a holophyletic group sensu Ashlock STAND: D va 42 Fig. 19. Diaprepocoris personata Hale: metaxyphus and ancilliary structures. Abbreviations; e, eye; 11p, lateral lobe of prothorax; 0, osteole of scent gland; P3, insertion of 3rd coxa; M, metaxyphus, scale line 1 mm. Hungerford (1948) split the Corixidae into six subfamilies: 1. Groups with large exposed scutellum .......... ee Diaprepocorinae and Micronectinae - Groups with scutellum hidden ................ 2 2. Hemielytral groove absent ..... Stenocorixinae = Hemielycralicroovespresentarre ee ee eee 3 3. Rostrum without transverse sulcations or nodal FUECO Ww ACEP PESI AP TEN Cymatiinae — Rostrum with transverse sulcations and nodal furrow ......... Corixinae and Heterocorixinae LANSBURY: Cuticular blades in Corixidae ' Figs. 20-23. Diaprepocoris pedderensis Knowles. - 20, 21, hind tarsi, scale line 1 mm; 22, middle leg, scale line mm; 23, hind femur and tibia, scale line 0. TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Popov (1986) recognised three extant subfami- lies viz. Corixinae, Diaprepocorinae and Micronec- tinae. Popov (1971) speculating on the affinities of Stenocorixa noted that the male genital capsule differed from all other corixids and that further study of the aberrant groups (Stenocorixa and Heterocorixa etc.) would be necesssary before the status of these groups was fully understood. Nachtigall (1974) comments ‘The rowing legs of recent water dwellers are considerably more flat- tened than those of fossil water dwellers or the legs of terrestrial relatives’. Within the Corixidae, both extant and fossil, there is little apparent distinc- tion. The fossil genus Gazimuria looks like a form of Diaprepocoris, Stenocorixa has explanate hind legs, Corixa moderately so, but there does not seem to be a steady progression throughout the corixid genera. As no corixids are remotely terrestrial, Nachtigal’s observations do not seem to be appli- cable. Immature stages Diaprepocoris barycephala 4th-5th instar: The front leg (fig. 25) closely resembles the adult. As in all immature corixids the front tibia-tarsus are fused, this character persists in the adult Diaprepocoris. The middle leg also closely resem- bles the adult form, but lacks the swimming hairs on the tibia-tarsus (fig. 26). The hind leg similar to the adult with prominent cuticular blades (figs. 12 & 27). The hind tarsus is one-segmented as in all known immature corixids. Immature stages of Stenocorixa were not avail- able for study. ACKNOWLEDGEMENTS The Leverhulme Foundation, London; CSIRO Division of Entomology, Canberra and the Austral- ian Biological Resource Study, Canberra are thanked for grants in aid towards a field-trip in 1979. Dr. W. D. Williams, Adelaide University, Dr. I. A. E. Bayly and Dr. P. S. Lake, Monash University sent me Diaprepocoris specimens. The SEM fig- ures were made by Mrs Barbara Luke, Zoology Department, Oxford University, courtesy of Pro- fessor D. S. Smith (Hope Professor). REFERENCES Ashlock, P. D., 1971. Monophyly and associated terms. — Systematic Zoology 20: 63-69. China, W. E., 1943. Some remarks on Walton's classifica- tion of the British Corixidae (Hemiptera). - Entomol- ogist's Monthly Magazine 79: 109-111. Horvath, G., 1927. Zoological results of the Swedish Expedition to Central Africa 1921, Insecta 25 (aquatic 44 = LE. ae a === = === III: | AN /; Mh SS / Ht Dil N Hal M I 7, / TE SF Fe —— = SS EZ ZEEE = SE II SZ = PI) hi N un Aly Di My N N Mh _ — Fig. 24. Diaprepocoris pedderensis Knowles, male dorsal habitus, scale line 1 mm. and semi-aquatic Hemiptera). - Arkiv för Zoologi 18A (31): 1-4. Hungerford, H. B., 1948. The Corixidae of the Western Hemisphere (Hemiptera). — Kansas University Science Bulletin 32 (25): 1-827. Jaczewski, T., 1928. Bemerkungen tiber die geographische Verbreitung der Corixiden. - Annales Musei Zoologici Polonici 7: 45-67. Kirkaldy, G. W., 1897. Aquatic Rhynchota: Descriptions and notes. No. 1. - Annals Magazine Natural History (6) 20: 52-60. LANSBURY: Cuticular blades in Corixidae ER GZ es I / AS i Mil! 7, ene A = — a — / \ | = = Il Sas \ SS m NT À = INN) TERN Ä | a / pd 4%, Figs. 25-27. Diaprepocoris barycephala Kirkaldy, immature. - 25, front leg, scale line 0.5 mm; 26, middle leg; 27, hind leg, scale line 0.5 mm. 45 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Leston, D., 1955. Taxonomy of the British Corixidae (Hem.). - Entomologist's Monthly Magazine 90: 57- 59. Lundblad, O., 1928. Studien iber schwedische Corixiden VII. - Entomologisk Tidskrift 49: 9-16, pls 1-3. Nachtigall, W., 1974. Locomotion: Mechanics and hydro- dynamics of swimming in aquatic insects. - In: The Physiology of Insecta, 2nd edition (edited M. Rock- stein): 382-432, 3, Academic Press New York. Parsons, M. C., 1976. Respiratory significance of the tho- racic and abdominal morphology of three Corixidae, Diaprepocoris, Micronecta and Hesperocorixa Heteroptera: Hydrocorisae. - Psyche, Journal of En- tomology, Camb. Mass. 83: 132-179. Poisson, R. & Jaczewski, T., 1928. Additional notes on the morphology of Stenocorixa protrusa Horvath (Heteroptera, Corixidae). - Annales Musei Zoologici Polonici 7: 115-119. Popov, Y. A., 1971. Historical development of Hemiptera Infraorder Nepomorpha (Heteroptera). - Trudy Pa- leontologicheskogo Instituta Akademiya Nauk SSSR, 129: 1-228 (in Russian). Popov, Y. A., 1986. Insects in the early Cretaceous eco- systems of West Mongolia. - Transactions Joint Soviet Mongolian Palaeontological Expedition 28: 50-83 (in Russian). 46 Schenke, G., 1965. Die Ruderbewegungen bei Corixa punctata Illig. (Cryptocerata). - Internationale Revue gesamten Hydrobiologie und Hydrographie 50: 73- 84. Schenke, G., 1966. Schwimmgeschwindigkeit, Schlagfre- quenze und Stevern bei Corixa punctata Illig. - Zoo- logischer Anzeiger 176: 5-12. Theiss, J., 1982. Generation and radiation of sound by stridulating water insects as exemplified by the corix- ids. - Behavioral Ecology and Sociobiology 10: 225- 235. Thorpe, W. H. & Crisp, D. J., 1949. Studies on Plastron Respiration IV. Plastron Respiration in the Coleo- ptera. - Journal Experimental Biology 26: 219-260. Walton, G. A. in Hutchinson, G. E., 1940. A revision of the Corixidae of India and adjacent regions. - Transac- tions Connecticut Academy of Arts & Science 33: 339- 476. Received: 20 March 1991 Accepted: 22 April 1991 NICO NIESER! & PINGPING CHEN? ‘Tiel, The Netherlands & “Organizing Committee XIXth International Congress Entomology, Beijing, P. R. China NAUCORIDAE, NEPIDAE AND NOTONECTIDAE, MAINLY FROM SULAWESI AND PULAU BUTON (INDONESIA) Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), I. Nieser, N. & P. Chen, 1991. Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heter- optera), I. - Tijdschrift voor Entomologie 134: 47-67, figs. 1-21. [ISSN 0040-7496]. Published 1 July 1991. Apart from new records for many species, mainly for Sulawesi, P. Buton and Sabah (N. Borneo), eight new species are described. Naucoridae: Aphelocheirus breviculus sp. n. (Sabah), A. robustus sp. n. (Sulawesi), Coptocatus stereos sp. n. (Sabah); Nepidae: Ranatra sulawesti sp. n. (Sulawesi); Notonectidae: Enithares caesaries sp. n., E. lansburyi sp. n., E. phenakismos sp. n. (all from Sulawesi) and E. skutalis sp. n. (P. Buton). Correspondence: Dr. N. Nieser, Htg. Eduardstraat 16, 4001 RG Tiel, The Netherlands. Key words. - Sulawesi; Buton; Sabah; Nepomorpha; keys; new species. During the last decades the study of Nepomor- pha and Gerromorpha of Indonesia and Malaysia is making some progress (see e.g. Lansbury 1964- 1985, La Rivers 1970, Polhemus 1986, Polhemus & Polhemus 1988). New expeditions to formerly un- explored areas and even rather occasional trips to already explored regions (see e.g. Nieser & Chen 1991), however, are still adding new species and new distributional data to our knowledge. As in many other groups, a high proportion of the spe- cies in this region seems to have limited ranges. In water bugs these restricted ranges are usually ap- parent in stream inhabiting species, whereas many of the widespread species occur in or on ponds in cultivated areas. The species dealt with in this paper were mainly collected by J. P. Duffels, J. van Tol and N. Nieser on several expeditions to the eastern part of Su- lawesi, the nearby island of Buton and Borneo. Keys to the Malesian genera are included. Keys to species of Sulawesi and Pulau Buton are given for most genera, but they should be used with extreme caution as the fauna of the region is still poorly known. Measurements are in millimeters and present the range or the (arithmetic) mean based on five randomly chosen specimens, or in case there are less than five, on all specimens available. The width of the head is measured across the eyes. MATERIAL A list of collecting localities of N. Nieser is presented in appendix 1. Sampling stations of J. P. Duffels and J. van Tol are partly connected with ‘Project Wallace’ and with the ‘1989 RMNH Expe- dition to Sulawesi’. Data on the latter expedition will extensively described in a separate paper (Van Tol et al. in prep.). All other sampling stations of both collectors, including those on Borneo, are summarized in appendix 2. In addition to these recent collections, miscel- laneous Malesian material in the RMNH collection has been included. Specimens collected by J. P. Duf- fels are deposited in ZMA, those by J. van Tol in RMNH, and those by N. Nieser in his own collection, unless stated otherwise. Deposition of material is specified only with rare or new species. The areas in Sulawesi used with the localities agree with the Indonesian provinces, viz. Sulawesi Utara for Northern Sulawesi (Celebes), Sulawesi Tengah for Central Sulawesi and Sulawesi Teng- gara for Southeastern Sulawesi. Pulau Buton is mentioned separately although it belongs adminis- tratively to Sulawesi Tenggara (see Whitten et al. 1988). Localities are in Indonesia, unless stated otherwise. Collections from which material has been stu- died (with abbreviations in brackets): Museum 47 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Zoologi Bogor, Bogor, Jawa (MBBJ); Bagian Pertan- ian, Universitas Haluoleo, Kendari, Sulawesi (BPUH); J. T. Polhemus collection (University of Colorado Museum), Englewood, Co., U. S. A. (yrPC); Snow Entomological Museum, University of Kansas, Lawrence, Kansas, U. S. A. (SEMC); N. Nieser collection, Tiel, The Netherlands (NC); Ox- ford University Museum, Oxford, United King- dom (OXUM); National Museum of Natural His- tory (Rijksmuseum van Natuurlijke Historie), Leiden, The Netherlands (RMNH); Zoölogisch Mu- seum, Afdeling Entomologie, Amsterdam, The Netherlands (ZMA). ACKNOWLEDGEMENTS Thanks are due to Dr J. P. Duffels (ZMA) and Mr. J. van Tol (RMNH) for putting specimens in their care to our disposal; to Dr I. Lansbury (OxUM) for discussing some Notonectidae and to Dr I Made Ebeneser (BPUH) for showing the senior author some interesting localities. SYSTEMATIC PART NAUCORIDAE Key to Indonesian and East Malaysian genera 1. Rostrum slender, longer than fore femora, reaching to middle coxae (Aphelocheirinae) AASTRA NET Aphelocheirus Westwood — Rostrum stout, very broad at base, much shor- ter than fore femora (Naucorinae) .......... 2 2. Rostrum inserted at anterior margin of head, labrum distinct and well developed ......... 3 - Rostrum inserted in a deep excavation, dis- tinctly posterior to the apical margin of head, labrum often greatly reduced (Cheirochelini) 3. Fore tarsi one-segmented, with one single very small claw (Naucorini) Naucoris Fabricius - Fore tarsi two-segmented, with two claws, which are often very inconspicuous (Laccoco- Tin) MINAS WOR TO NET MAENE de 4 4. Labrum more or less equilaterally triangular with antacuteltiprs sas eee Laccocoris Stal - Labrum with sides shorter than base and tip broadly rounded ............... Heleocoris Stàl 5. Head ventrally with well developed grooves in which litthe antennaes ER PURE REA PROTO 07 L'EAU Coptocatus Montandon = Antennal oroovesfabsentan. MN 6 6. Ventral margin of prothorax with a conspicu- ous pit covered with amembrane or specialized pilosity (static sense organ), body elongate, 48 connexival angles blunt, not spinose, conse- quently lateral edge of abdomen more or less CONTINUOUSEN RE CT 7 - Static sense organ on ventral margin of pro- thorax absent, body oval, connexival angles sharp, usually spinose, consequently lateral edge of abdomen distinctly serrate .......... 9) 7. Static sense organ slightly anterior to midway of lateral margin of prothorax, ventral keel of head strongly developed with an anterior spine-like tip, which is at least as long as first rostral segment ......... Idiocarus Montandon - Static sense organ near or at antero-lateral an- gle of prothorax, ventral keel of head normal, without a prominent anterior spine ......... 8 8. Static sense organ rounded, placed at extreme antero-lateralangle of prothorax, which is cup- shaped; anteclypeus short, broadly and evenly LOUNGEC A Sete Nesocricos La Rivers — Static sense organ elongate, placed half to three-quarters of its length caudad to antero- lateral angles of prothorax; anteclypeus quite long with sinuate margins producing a medio- ANLELIOLIODER EP EEE Tanycricos La Rivers 9. Mesotibia distally with five or six decreasingly sized, transverse rows of dense, short spines en RN ne Cavocoris La Rivers - Mesotibia with at most two of such rows of Spines 10.121092. ADN BORO AGREE ACER SET AUS 10 10. Anteclypeus prolonged, overhanging part be- ing half to three-quarters the length of labral baser dE AEEA EM Warisia La Rivers — Anteclypeus short, overhang at most one third of the length of labral base ........... JE 11. Abdominal venter showing only four segments ERICE. WI ANA IA, Quadricoris La Rivers - Abdominal venter showing the usual five seg- mentst SINE FE E ROTA RR, 12 12. Larger species, length over 7 mm; abdomen asymmetrical ........... Sagocoris Montandon - Smaller species, length slightly less than 7 mm, AbdoMENisyimmetrica MORE REI ROLL San Aptinocoris Montandon Remarks. - Except for Sagocoris, which has also one representative in the Philippines, the last mentioned eight genera are restricted to New Gui- nea. In the species of Naucoridae described below, the first abdominal segment is not visible. The numbers used are the morphological homologous numbers, thus ‘tergites 2-8° are visible as tergites 1-7. Aphelocheirus Westwood, 1833 Southeast Asiatic species of the genus were re- cently revised by Polhemus & Polhemus (1988). About 18 species are known from Malesia, includ- ing the Philippines, but more species are to be discovered. Species of Aphelochetrus are easily overlooked by the casual collector as most burrow in the bottom of streams. Thanks to their plastron respiration they do not need to surface to refill their air store. Key to Aphelocheirus of Sulawesi 1. Lateroposterior spines of connexiva 4-5 short and broad; body length of male 8.9-9.1 mm, of female 8.5-8.8 mm (Sulawesi Selatan) ......... MERE Te Rue. à A. celebensis Polh. & Polh. — Lateroposterior spines of connexiva 4-5 rather long and acute; body length of male c. 10 mm, oftfemalesmoresthan 92mm. rer een. 2 . 2. Male: Left paramere with a broad hooked apex (fig. 5). Female: Genital operculum short and broad with convex caudo-lateral angles (fig. 3) (Sulawesi Tenggara) ...... A. robustus sp. n. — Male: Left paramere with a slender curved ap- ical part. Female: Genital operculum rather long and slender with apical lateral margins very slightly concave (Sulawesi Tengah) ...... LIA o EE ALU GF A. lorelindu Polh. & Polh. Aphelocheirus lorelindu Polhemus & Polhemus Aphelocheirus lorelindu Polhemus & Polhemus, 1988: 190-192, figs. 18-21, 164. Material. - Sulawesi Tengah: Palu, 50 km SE of: Lore Lindu NP, Sopu river near Dongi Dongi, 950 m, SJ 86, 8 Dec 1985, J. van Tol, 3 9, (RMNH). Distribution. - Known only from the Lore Lindu N. P. in Central Sulawesi, and surrounding areas. Aphelocheirus breviculus sp. n. (figs. 1-2) Type material. - Holotype 9: Malaysia: S. Sabah: 105 km S of Beaufort, Long Pasia area, Sungai Ritan, 4°24'’N 115° 42’E, 1160 m, undisturbed evergreen tropical rain- forest, 9.IV.1987, J. van Tol & J. Huisman (RMNH). - Paratypes: 2 ® with same data (RMNH, NC). Description Brachypterous female. - Rather small species, form elongate oval, length 7.7-8.1 mm, width of head 1.8-1.9 mm, maximum width across abdomen 4.9-5.1 mm. Colour greyish brown, head on dorsal side yellow to light brown, posteriorly darker, with black eyes; pronotum, hemielytra and tergites greyish to brown, margins and scutellum paler; legs and ven- NIESER & CHEN: Malesian Heteroptera ter yellowish to light brown, central part of tho- racic and abdominal sternites greyish, pruinose, with sparse golden pubescence. Head. Shining, coarsely punctate, produced ahead of eyes for distance equal to 0.5 the length of an eye; eyes twice as long as wide, outer margins sinuate, due to weakly developed antero-lateral flange; anterior / posterior interoculus = 1.3/1.0 mm. Thorax. Pronotum shining, roughly punctate, antero-medially rugose; very sparsely set with in- conspicuous short fine pale setae, width / length 3.5; lateral margins bearing about twelve stout minute erect setae. Scutellum shining, rugose, width / length 2.5, lateral margin hardly sinuate, shallow transverse sulcus present along anterior margin (to fit the posterior margin of pronotum). Hemielytra not touching medially, leaving an area 0.4 mm wide of metanotum uncovered; posteriorly not reaching the hind margin of abdominal tergite 2, surface punctate-rugose, claval area shining, dull in remainder, clavus not delineated; embolar mar- gin evenly curving anteriorly, its posterior angle strongly produced laterally (fig. 1). Abdomen. Dull, rugose with narrow shining margins which bear 5-7 minute erect setae in small indentations, posterior indentation of each seg- ment bearing two or three of these minute setae and a few longer, easily detachable, setae. Tergites 2-8 exposed, bearing very few short thin pale re- cumbent setae, paired glandular openings present, but indistinct on posterior margin of tergite 3. Posterolateral angles of tergites 3-7 bluntly and broadly spinose, more distinctly produced poste- riorly. Ventral side. Length of antennal segments 1-4 is 0.09, 0.23, 0.23 and 0.42 mm. Labrum shining, anterior margin evenly rounded, rostrum shining, length 3.2-3.4 mm, reaching middle trochanters. Prosternum with weak median carina, propleura with inner projections notched; mesosternum pruinose with a median carina, tumescent poste- romedially; metasternum pruinose, metaxiphus small, apically pointed. Abdomen pruinose, ster- nites 3 and 4 bearing rounded projections medially on posterior margins, posterior margins of ster- nites 4 and 5 lacking rearwardly directed stout setae medially; genital operculum with posterolateral angles developed, resulting in a sinuate posterior margin; narrow tufts of long setae at lateral mar- gins and intermedially in apical third (fig. 2). Legs. Set with fine gold setae, fore femur, tibia and tarsi with thick hair pads on inner surface. Fore, middle and hind coxae with combs of long light brown setae on interior margins. Middle leg, trochanter, femur, tibia and tarsi with thick yellow hair pads on ventral faces; femur sparsely set with 49 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 1-5. Aphelocheirus species. - 1, Apex of em- bolium of A. minor (left) and A. breviculus (right); 2, Genital operculum of 9 A. breviculus, 3, idem, A. robustus; 4, A. robustus, apex of abdomen of bra- chypterous 4, dorsal view; 5, A. robustus, parameres, ventral view. Scales: 1 mm (1-3), 2 mm (4), 0.5 mm ©): short stout reddish spines, bearing 6-8 long erect setae on posterior margin; tibia rather densely set with short stout reddish spines, bearing about four long setae on anterior margin and one row of red spines apically. Hind leg, femur and tibia sparsely set with short stout reddish spines, tibia also with an apical transverse row of reddish spines, tibia and tarsi bearing long swimming hairs on ventral sur- face. Claws of all tarsi shining, curved, with darker, reddish brown apices. Comparative notes This species runs to A. lahu Polhemus & Polhe- mus and A. minor Polhemus & Polhemus in the key of Polhemus & Polhemus (1988). It shares with A. minor, which is also from Sabah, abdominal segments 3 and 4 bearing rounded projections me- dially on the posterior margins. A. breviculus, how- ever, is one millimeter longer on average, has the angle of the embolium more produced (fig. 1), and 50 the ratio width / length of the pronotum greater than in A. minor, viz. 3.6 and 3.2 respectively. The eyes of A. minor are narrower, about three times as long as wide, while A. breviculus has its eyes twice as long as wide. Etymology Breviculus, a Latin adjective, meaning rather short, refers to the small size of the species. Aphelocheirus robustus sp. n. (figs. 3-5) Type material. - Holotype @ (brachypterous): Su- lawesi Tenggara, K. Kolaka. Sungai Mowewe, 28. II. 1989, leg. N. Nieser, N8922 (RMNH). — Paratypes: 9 @ 14 Q (incl. 1 labelled allotype) brachypterous, 16 @ 7 9 ma- cropterous, same data as holotype, deposited as follows 3 & 3 Q brach., 3 6 3 Q macr. (NC); 1 9 brach., 1 & macr. (MBBJ); 1 6 2 2 brach., 2 & 1 9 macr. (JTPC); 1 Q brach., 2 & macr. (SEMC); 1 & macr., 1 Q brach. (OxUM); 1 9 brach., 1 & 1 9 macr. (RMNH); 1 @ 1 Q brach, 1619 macr. (ZMA). Additional material. - 18 larvae IV/V, with same data as holotype. Description Brachypterous male. - Large species for this ge- nus, form elongate oval, length 9.8-10.5 mm; width men) 5.9-6.2 mm. Colour. Somewhat variable. Dorsal side: head yellow with black eyes, pronotum yellow with an- terior and posterior transverse dull brown band, scutellum yellow (most specimens) to brown, he- mielytra brown with variable part posteriorly yel- low, abdominal tergites dull brown with yellow posterolateral angles, genital segments yellow. Ventral side: head, including antennae yellow, ros- trum pale brown, sternites brown with yellow lat- eral margins, legs and genital segments yellow. Head. Shining, finely rugose, produced ahead of eyes for distance equal to 0.4 X the length of an eye; eyes length / width = 2.4-2.5, outer margin sinuate due to weakly developed anterolateral flange; an- terior / posterior interoculus = 1.35-1.40. Thorax. Pronotum rugose, sparsely set with short fine pale setae laterally, apparently glabrous medially, width / length = 3.7-4.0; lateral margins bearing c. 10 stout minute erect setae. Scutellum rugose, appearing glabrous, width / length = 2.3- 2.5, lateral margin hardly sinuate, shallow trans- verse sulcus present along anterior margin (to fit the posterior margin of pronotum). Hemielytra touching medially, reaching posteriorly to or just beyond base of abdominal tergite 3, surface rugose, set with sparse fine pale setae, embolar margin evenly curving anteriorly, bluntly terminated pos- teriorly in most specimens. Abdomen. Weakly rugose, set with fine pale se- tae, tergites 2-8 exposed, paired glandular open- ings present medially on posterior margin of ter- gite 3, posterior margin of tergite 5 asymmetrical, with a medioposterior hump-like projection, which is delimited more clearly on the left than on the right (fig. 4), lateral margin of all segments with short stout setae, posterolateral angles of all tergites 3-7 spinose, with a few long pale setae at base of spinose angles. Ventral surface. Length of antennal segments 1 to 4: 0.10, 0.20, 0.35 and 0.45 mm; rostrum gla- brous, length 3.2-3.4 mm, reaching to middle coxae. Labrum yellow, shining, anterior margin evenly rounded. Prosternum with weak median carina, propleura with inner projections notched; mesosternum pruinose with a median carina, tu- mescent posteromedially; metasternum pruinose, NIESER & CHEN: Malesian Heteroptera metaxiphus small and narrowly pointed. Abdomen pruinose, posterior margins of sternites 4 and 5 with six rearwardly directed stout setae medially; genital segments glabrous, shining. Legs. Set with fine golden setae, fore femur, tibia and tarsi with thick hair pads on inner surface, claws well developed and curved in most specimens but in several short, stunted and not curved. Fore, middle and hind coxae with combs of long light brown setae on anterior margins. Middle legs, tro- chanter, femur, tibia and tarsi with thick yellow hair pads on ventral faces; femur sparsely set with short stout reddish spines bearing 6-8 long erect setae ON posterior margin; tibia sparsely set with short stout reddish spines, bearing three long setae basally on anterior margin and one row of red spines apically. Hind leg, femur sparsely set with short stout reddish spines, tibia with reddish spines along interior margin, tibia and tarsi bearing long silvery swimming hairs on ventral surface, claws shining, curved. In several specimens middle and hind claws and sometimes tarsi stunted. Male genitalia. Right paramere longest with long golden pubescence on right margin, apex rounded, left paramere hooked with thick golden pilosity on left margin. (fig. 5). Brachypterous female. - Length 9.5-10.1 mm; width of head 2.2-2.3 mm; maximum width 5.8-6.0 mm. Subgenital plate triangular with narrow lat- eral earlike structure (fig. 3), longer lateral setae and shorter posterior setae spread out, not in nar- row tufts. Macropterous form. - As brachypterous, except length & 10.6-11.0 mm, 2 10.0-10.6 mm. Prono- tum well developed, subequal to maximum width of abdomen; scutellum larger and somewhat in- flated; mesosternum inflated with broad rounded carina medially; hemielytra well developed (mem- brane broken off in some females), blackish in most fully mature specimens, leaving lateral part of connexiva uncovered; embolium with a blunt but distinct angle midway. Comparative notes This species runs to A. lorelindu Polhemus & Polhemus in the key to brachypterous specimens of Polhemus & Polhemus (1988); both species are very similar in general shape and size. The female subgenital plate of A. robustus differs from larger Southeast Asiatic Aphelochetrus by its small lateral ‘ears’. The right paramere is narrower at base than A. lorelindu and A. celebensis Polhemus & Polhe- mus, its closest relatives. See also the key to the Sulawesi species. 51 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 | Figs. 6-7. Coptocatus stereos. - 6, Left paramere n. Scale 0.5 mm; 7, Tibiotarsus and anterior margin of femur of fore leg. Scale 1 mm. Etymology Robustus, a Latin adjective, refers to the large size of the species within the genus. Remarks The type locality is a lowland stream, 2.5 to 10 m wide, flowing through woodland with fields and small settlements close by. Aphelocheirus was found at shallow, partly shaded places, with mod- erate current and coarse sand bottom. Benthic fauna, present in rather low density, included mainly Ephemeroptera, with a few Odonata and Trichoptera. The stunted claws and tarsi may be the results of wear and tear in older specimens, but this pheno- menon is mentioned in the description, since quite a few specimens are strikingly affected. Coptocatus Montandon, 1909 The genus Coptocatus was recently revised by Polhemus (1986). Including the species described below, four species are known from the eastern states of Malaysia (Sabah and Sarawak). 52 Coptocatus stereos sp. n. (figs. 6-7) Type material. - Holotype & (brachypterous): Malay- sia: S. Sabah. 105 km S of Beaufort: Long Pasia area, Sg. Maga near confluence Sg. Pasia. 4°26'N 115°40’E, 1210 m asl., larger fast running stream in untouched lower montane evergreen rain forest, large boulders, rapids. 3 Apr. 1987. Leg. J. van Tol (RMNH). Additional material. - 3 larvae instar V and 1 instar III / IV, same data as holotype (RMNH). Description Holotype @. - Large, robust, highly streamlined and dorsoventrally flattened; general colour me- dium brown with yellowish spots. Length 20.4 mm, width of head 5.4 mm; width of pronotum 10.6 mm, maximal width 12.2 mm. Head. Width / length 5.4 / 4.2 mm. Anteclypeus greatly produced, apex evenly rounded, projecting beyond rostrum for a distance greater than exposed rostral length when viewed laterally; eyes blackish, convex, roughly rectangular, length / width 1.8 / 1.2 mm, raised above level of vertex and separated from it by wide shallow sulci, lateral margins bear- ing a weakly developed flange; anterior / posterior interoculus 3.2 / 3.2 mm; vertex greatly produced posteriorly, extending rearwardly about one half the length of an eye; anteriorly with a pair of broad shallow sublateral depressions, tapering poste- riorly. Thorax. Pronotum width / median length 10.4 / 3.0 mm; lateral margins with narrow, raised darker and slightly crenulate ridge, posterolateral angles strongly produced, rounded. Scutellum width / length 5.0 / 2.8 mm, anterior margin strongly reflexed downward. Hemielytra brachy- pterous, extending to posterior margin of abdom- inal tergite 3 only, tips broadly rounded, surface set with fine yellowish granular microstructure (also present on other parts of body, although far less dense and less distinct), embolium anteriorly de- fined by a broad carina, explanate, lateral margin irregularly and shallowly crenulate, set with long recumbent light brown setae; hemielytral commis- sure with a small projecting tooth on left hemie- lytron distally, fitting into corresponding indenta- tion of right hemielytron. Abdomen. Medio- and latero-caudal parts of ter- gite 3 and all of tergites 4-8 exposed. Lateral mar- gins with narrow ridges and a dense row of long light brown setae lying flat on tergites. Medially some patches of shorter setae at base of tergites, especially the lobes of tergite 8. Lateroposterior angles indented with blunt projections and a tuft of hairs in the indentation. Ventral surface. Sternites pruinose. Gula and part of prosternum anterior to fore coxae with a blunt carina, posterior part of prosternum without carina. Mesosternum with a shallow median groove and a distinct xiphus on posterior margin. Metasternum anteriorly with a strongly dorsally declivent carina, posterior part flat with a short broad blunt xiphus. Abdominal sternites with large stigmata halfway of lateral third; sternites 4, 7 and 8 with small tufts of short golden hairs medially, sternite six medially produced, sternites 5-8 me- dially glabrous. Legs. Fore leg with massive femur, length 6.5 mm, width 4.3 mm; suture between tibia and tarsus shallow, claw short, blunt, hardly differentiated; anterior face of tibiotarsus and femur with golden pubescence distally becoming thicker and longer and extending onto trochanter; anterior margin of femur with a median hump and a subapical tooth (fig. 7). Posterior margin of all femora with a fringe of long hairs. Middle leg and hind femur without spines. Middle and hind tibia ventrally with a thick pad of golden brown pilosity (looks like a adhesive pad; already fully developed in larva V, and partly so in larva III/IV). Middle tibia broa- dened apically. Hind tibia with short spines along inner and outer faces, inner (posterior) faces with a dense fringe of long swimming hairs. Genitalia. Parameres symmetrical, bilobed, outer lobe swollen (fig. 6). Aedeagus symmetrical, basally and centrally chitinized, apically and later- ally fleshy. Comparative notes The same size and similar anterior margin of fore femur as C. kinabalu D. Polhemus, also from Sabah, which has, however, the inner lobe of para- mere much shorter and the lateroposterior angles of abdominal segments not distinctly indented. The other two species in the genus known from Sabah and Sarawak do not exceed 15 mm in length. Etymology Stereos, a greek adjective, meaning: strong, solid, cruel, refers to the size and solid build of the species. Remark The holotype has a case, probably of a chiro- nomid larva on the left hemielytron against the inner embolar ridge. Laccocoris Stal, 1856 Seven species are known from Malesia, but none has been reported from Sulawesi (La Rivers 1971). The same is true for the genus Heleocoris Stal with six Malesian species. The last revision of both genera is Montandon (1897). NIESER & CHEN: Malesian Heteroptera Laccocoris staudingeri Montandon Laccocoris staudingeri Montandon, 1897: 440-442. Material. - Malaysia. S. Sabah, Long Pasia area, Sg. Ritan, 4°24'N 115°42’E, 1160 m asl. Leg. J. Huisman & J. van Tol, 16 1 9 (RMNH). Distribution. - Brunei and Sabah. Remarks. - L. staudingeri differs from related species in northern Borneo, L. horvathi Mont. and L. maat La Rivers (La Rivers 1970) by having the interoculus at halfway level slightly more than twice as wide as an eye. Naucoris Fabricius, 1775 There are six Malesian species, of which one occurs in Sulawesi. A revision of the genus is not available. Naucoris scutellaris Stal Naucoris scutellaris Stal, 1859: 266. - Lundblad 1933a: 63-67, fig. 19 (redescription). Material. - Sulawesi Tenggara: Kendari, N8904, 1 8; Mowewe, N8922A, 4 & 2 9; 20 km S Pomalaa, N8928 (NC). Remarks. - A widespread species, occurring from India trough Sri Lanka and Thailand to Jawa and Sulawesi. The femur is strongly expanded at its inner apical two-thirds, giving the inner margin a deeply dented appearance. Other congeneric spe- cies have the inner margin of fore femur more or less straight. NEPIDAE Key to Indonesian and Eastern Malaysian genera 1. Flattened species, parasternites of abdomen visible, head distinctly narrower than the pro- notum and partly enclosed by its anterolateral ANOTESI(INE PINA) RE ERRE E ere 2 - Subcylindrical species, parasternites not vis- ible, head free from pronotum and both of subequal width (Ranatrinae) ................. 3 2. Respiratory siphon longer than the inner mar- gin of the hemielytra ....... Laccotrephes Stàl - Respiratory siphon not more than half as long as the distance between apex of scutellum and the membrane along inner margin of hemi- elytra (only one very rare species, T. breddini Montandon, from Northern Borneo) .......... SPARI STIRIA Telmatotrephes Stàl TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 3. Eyes in lateral view globose, not reflexed downwards and not obscuring ventral margin OI NEA ET na neer Ranatra Fabricius — Eyes in lateral view reflexed downwards, obs- curing ventral margin of head .................. PRAIA AI BAE Cercotmetus Amyot & Serville Laccotrephes Stal, 1865 About eight species recorded from Malesia. A generic revision is not available and the status of many species is uncertain, which makes some re- cords unreliable. Three species are known from Sulawesi and Pulau Buton. Key to Laccotrephes of Sulawesi and Buton 1. Body length over 35 mm, usually 40 mm or MOL ay eet Jenssen dek L. robustus Stal - Body length less than 35 mm ................ 2 2. Respiratory siphon as long as or somewhat shorter than body ....... L. occultus Lundblad - Respiratory siphon distinctly longer than body MCO ERI E ee Si L. tristis (Stal) Laccotrephes occultus Lundblad Laccotrephes occultus Lundblad 1933a: 23-26, fig. 2, tab. 14 (1) [nom. nov. for L. fuscus auctores, nec. L., redes- cription]. Material. - P. Buton: 16 km E of Baubau, N8944, 1 & (NC). Maluku: Bacan, Wayauna, c. 50 m alt., logged forest, hand coll., 23-26 Jun 1985, leg. J. Huijbregts, 2 9 (RMNH). Distribution. — Sumatera, Borneo, Buton, and Maluku (Moluccas). Laccotrephes robustus Stàl Laccotrephes robustus Stal, 1870: 706. - Lundblad, 1933a: 26-27, fig. 3 [redescription]. Material. — Jawa: Preanger, 1 @ 1 Q; Paluabuan, 3.X1.1911, 1 g. Sumba: Kananggar, 700 m, V.1925, 1 à; Mao Marroe, 450 m, V.1925, leg. Dammerman, 1 9. Malaysia (Sabah): Danum valley, 4°48’N 117°48’E, 220 m streamlet, 23.III.1987, leg. J. Huisman, 1 9 (all RMNH). Distribution. - Widespread species: Indochina and Taiwan, Philippines, Malacca, Sumatera, Jawa, Borneo, Sulawesi, Sumba and Bali. Laccotrephes tristis (Stàl) Nepa tristis Stàl, 1854: 11. Laccotrephes tristis; Hale, 1924: 504-507, pl. 34 (figs. 1- 3), pl. 35 (figs. 2, 4, 6), pl. 36 (fig. 18); Lundblad 1933a: 25; Lansbury 1967: 644-646 [description of male ge- nitalia]. 54 Material. - Sulawesi Utara: Dumoga Bone N. P., To- raut, c. 200 m, multistr. evergreen forest, creek, 23.V.1985, leg. J. Huijbregts, 1 Q (RMNH). Distribution. — Previously only known from Australia and New Guinea. Remarks. - Identification of this specimen is doubtful. It differs from typical L. occultus by its slightly greater size (34 vs. 31-32 mm), its nar- rower appearance anteriorly (only visible under direct comparison), and its much longer respira- tory siphon (40 vs. 30 mm). Cercotmetus Amyot & Serville In the excellent revision of the genus by Lans- bury (1973), six species are recorded from Malesia, including one from Sulawesi. Most species of this genus are seldomly collected and then usually only one specimen at a time. Consequently, distribu- tional patterns of most species are poorly known. Key to Malesian species of Cercotmetus Ea from Lansbury 1973) . Vertex raised between the eyes, but without a distinctituberclen sare EIER N Sr 2 — Vertex with a distinct, broadly and tooth-like tubercle rare Rene RUE 4 2. Less than 43 mm long, middle femora clearly shorter than prothorax (widespread species: Sumatera, Jawa, Sarawak) aan aaa cs C. brevipes Montandon - More than 47 mm long, middle femora longer thanyprothorax SERIA AE RR REN. 3) 3. Eyes relatively small, width of vertex halfway the inner margin of eyes about twice the width of an eye at that level, vertex with a complete eireletiof/hairst(Sulawes ep mr IB AINSI, 43 C. robustus Montandon — Eyes larger, width of vertex halfway the inner margin of eyes subequal to the width of an eye, hairs on vertex few, not forming a complete circlet posteriorly (Irian Jaya) ................... TS IRE, PH NO OSG C. dissidens Montandon 4. Distal third of tergite 6 raised and often very hairy, apex of paramere sinuate, female oper- culum not or hardly reaching beyond tergite 6 (widespread Sumatera)! war rue eee tae Hen SSR: OMAN AUS C. compositus Montandon - Distal third of tergite 6 not distinctly raised and never hairy, apex of paramere evenly curved, female operculum reaching beyond ter- gite 6 for about one third of its length ...... 5 5. Along median line length of anterior lobe (an- terior to transverse grooves) of pronotum less than three times the length of posterior lobe (widespread, Sumatra, Jawa, Borneo) .......... ae ee eee C. asiaticus Amyot & Serville - Along median line length of anterior lobe more than three times the length of posterior lobei(Kalimantan, Sarawak) 2... ES C. astaticus var. longicollis Montandon Cercotmetus robustus Montandon Cercotmetus robustus Montandon, 1911: 92-93. — Lans- bury 1973: 95-97, figs. 36-42 [redescription]. Material. — Sulawesi Tenggara: N8922A, 2 Q (NC). Remarks. — Apparently a rare species, only known from the female holotype from Central Sulawesi (‘Posso See’ = Danau Poso) so far. Several small larvae, possibly of this species, were observed at locality N8931 (see appendix). Both larvae and adults are far better swimmers than Ranatra, they seem to use their fore legs for swimming. Ranatra Fabricius, 1790 About twelve species recorded from Malesia, of which three have been reported from Sulawesi (one with two subspecies). The Oriental species have been revised by Lansbury (1972). In the fol- lowing key to the Sulawesi species, also two unre- corded but widespread species that might turn up in Sulawesi, were included. Key to Ranatra of Sulawesi (Adapted from Lansbury 1972). 1. Head with prominent tubercle between eyes, metasternum emarginate (fig. 9) ............ 2 - Head at most only slightly raised between eyes, metasternum not emarginate ................ 3 2. Length of respiratory siphon subequal to length of body (Sulawesi Tenggara) ........... PRE R. sulawesti sp. n. - Length of siphon one-third of body length (not recorded from Sulawesi, but a widespread spe- cies, Indochina, Sumatera, Jawa, Nusa Teng- PALA) Ae Eg R. parmata Mayr 3. Width of fore femora measured from dorsal margin to apex of large tooth about the same as widest part of femora proximally ........ 4 - Width of fore femora measured from dorsal margin to apex of larger tooth clearly greater than widest part of fore femur proximally (en- demic to Sulawesi) .... R. malayana Lundblad 4. Larger tooth of fore femora more or less equi- distant between ends of femora (not recorded from Sulawesi, but a widespread species: India, Ceylon, Indochina, Sumatera, Jawa) ............ Re el R. varipes Stàl — Larger tooth of fore femora clearly nearer to distallendo Memo ne MEET 3 NIESER & CHEN: Malesian Heteroptera 5. Third episternum with two clumps of small hairs (on each side) (endemic to Sulawesi Sel- atan) rer R. longipes celebensis Lansbury - Third episternum without clumps of small hairs (Indonesia, incl. Sulawesi Tenggara, E. Malaysia) no R. longipes longipes Ranatra longipes longipes Stàl Ranatra longipes Stàl, 1861: 203. Ranatra longipes longipes. - Lansbury 1972: 332-334, figs. 220-229 [redescription]. Material. - Sulawesi Tenggara: N8901, 3 & 3 9; N8909, 3 6; N8910, 2 6; N8917, 2 6; N8919, 1 & 3 9; N8930, 19. Distribution. - Known from Sumatera, Borneo (Brunei, Sabah), Jawa, Bali and Sulawesi. Remarks. - Our specimens were compared with a series from Bali, of which several were identified by Lansbury. The specimens mentioned above do not show any of the diagnostic characters charac- terizing subspecies celebensis (Lansbury 1972). Ranatra malayana Lundblad Ranatra birdi var. malayana Lundblad, 1933a: 40, fig. IA. Ranatra malayana. - Lansbury 1972: 319-321, figs. 142- 152. Material. - Sulawesi Tenggara: N8917, 1 @ 1 9 (NC). Remarks. - Until now only known by three spe- cimens from Sulawesi Selatan. Respiratory siphon, when folded back over dorsum, reaching halfway head in the male, and anterior margin of pronotum in female. Ranatra sulawesii sp. n. (figs. 8-11) Type material. - Holotype &: Sulawesi Tenggara, K. Kendari, pond with Nymphaea at Wawonggole, 20.11.1989, N8902, leg. N. Nieser (RMNH). - 10 6 79 (including one labelled allotype), same data as holotype, deposited as follows: 1 Q (RMNH); 3 6 2 Q (NC), 16 1 9 (MBBJ); 1 & (BPUH); 1& 1 9 (TPC); 1 6 1 9 (SEMC); 1 6 (OxUM). Description Adult. - Measurements. Length & 34-38.3 mm; Q 35-39.5 mm; length of siphon @ 39.0-41.7 mm, Q 35-45 mm; width of head @ 3.2-3.3 mm, Q 3.2- 3.5 mm; humeral width of pronotum & 3.5-3.9 mm; Q 3.6-4.2 mm. Colour. Dark brown, legs paler with indistinct annulation. 55 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 8-11. Ranatra sulawe- si. - 8, antenna; 9, metaster- num, 10, genital capsule of male, lateral view; 11, apex of paramere. Scales: 0.5 mm (8), 1 mm (9-10), 0.25 mm (11). Head with small, but distinct nodule on vertex; eyes slightly wider than width of interoculus; lora strongly developed, higher than clypeus, which has two small nodules apically; second segment of an- tennae with long, finger-like projection, subequal to length of segment 3 (fig. 8). Thorax. Prothorax in lateral view about 1.5 X as long as fore coxa, about 1.1 X as long as fore femur; anterior lobe twice as long as posterior lobe in median line; anterior margin not conspicuously raised; humeral width / anterior width 1.4; pro- notal grooves well developed; posterior lobe with- out tubercles, ventrally with a broad concavity an- teriorly at insertion of coxae, caused by produced anteroventral angles, remainder of prothorax flat with a vague carina in anterior one-third. Scutel- lum about 1.5 X longer than wide, with a shallow broad transverse depression in apical third. Meso- sternum with anterior margin raised, posterior margin of metasternum deeply incised (fig. 9); space between middle and hind coxae subequal. Legs. Fore femur with one median tooth at apical 0.4 of femoral length, no secondary tooth distally. Length of middle and hind femur subequal. Middle 56 tibia shorter, hind tibia longer than femur; apical % of middle and hind tibia with hair fringe. Hind femur reaching about % along last abdominal ster- nite in both sexes. Male genitalia. See figs. 10, 11. Female. - Genital operculum not reaching the base of respiratory siphon. Comparative notes Based on the tuberculate vertex and emarginate metasternum this species can be included in the R. gracilis group (Lansbury 1972). R. sulawesu can be distinguished by (a) siphon longer than the body length, (b) distance between the middle and the hind coxae subequal, and (c) finger-like projection of antennal segment 2 long. In the key by Lansbury (1972) it runs to R. parmata Mayr, from which it differs by its long siphon, and the shape of the pa- ramere, which is apically more swollen in R. parmata. Etymology Sulawesi, a noun in genitive case, refers to the type locality. Remark The type locality is a small pond at the edge of marshy fields with some water lilies (Nymphaea sp.), and abundant marginal vegetation flattened and hanging into the water. NOTONECTIDAE Key to Indonesian and Eastern Malaysian genera 1. Hemielytral commissure anteriorly without a definite hair-lined pit (Notonectinae) ...... 2 — Hemielytral commissure anteriorly with a def- inite hair-lined pit (Anisopinae) ............... nits. Mertens berte den Anisops Spinola 2. Middle femur with an anteapical pointed pro- tuberance Ara ere Enithares Spinola - Middle femur without such a protuberance 3 3. Eyes contiguous or forming an ocular commis- sure near posterior border of head ............. ee etudes Nychia Stal - Eyes widely spaced near posterior border of head fm mortes Aphelonecta Lansbury Anisops Spinola, 1840 Due to insufficient knowledge of the fauna, a key to the regional species of the genus Anisops Spinola cannot be presented. The reader is refer- red to Brooks’ (1951) revision and various papers by Lansbury (1964, 1965, 1978). Several species, e.g. A. breddini, A. nasuta and A. paracrinita, can be found in village ponds and rice fields. Such species tend to be very widespread in Southeast Asia. Anisops batillifrons Lundblad Anisops batillifrons Lundblad, 1933: 463-464, fig. 8. - Brooks 1951: 420-423, figs. 71, 101. Material. - Malaysia, Sabah: 16 km NE Tenom: Agri- cultural Research Station, Sg. Segalan, 115°59’E 5°12’N, a. Ll, 23.XI.1987, 1 & 1 Q, leg. J. Huisman & R. de Jong (RMNH). Distribution. — A widespread species: India through Indochina to Hainan, Taiwan, Iriomote and the Philippines; apparently mainly in subtrop- ical regions. Anisops biroi Brooks Anisops biröi Brooks, 1951: 454-456, fig. 49. - Lansbury 1978: 111, figs. 37-39. NIESER & CHEN: Malesian Heteroptera Material. - Sulawesi Tengah: 50 km SE of Palu: Lore Lindu N.P., Sopu valley near Dongi Dongi, 6 Dec 1985 (sample B), leg. J. van Tol & J. Krikken 3 & 2 9 (RMNH). Pulau Buton: N8939, 10 @ 4 Q (NC). Maluku: Bacan, Wayauna, alt. 50 m, sec. growth, eutrophic pool, 5-7 July 1985, HH382, leg. J. Huijbregts, 1 @ 5 ® (RMNH). Remarks. - Identification of this species some- what doubtful. The rather short rostral prong sug- gests the very similar A. rigoensis. None of the present specimens, however, has a carina on the posterior part of the vertex, which is reported to be characteristic for A. rigoensis (Lansbury 1978). Both species were only known from New Guinea up to now. Anisops breddini Kirkaldy Anisops breddini Kirkaldy, 1901a: 5-6. - Brooks 1951: 439-441, figs. 78, 99; Leong 1962 [life cycle]; Lansbury 1965: 58. Material. - Sulawesi Tenggara: N8901, 22 @ 27 9, 5 larvae; N8917, 1 6 1 9; N8918, 1 4 19; N8931, 1 6 1 ©. Pulau Buton, N8938, 5 & 22 9. Distribution. - Widespread species: India and Sri Lanka through Indochina to Jawa, Sulawesi and Buton. Anisops nasuta Fieber Anisops nasuta Fieber, 1851: 484-485. - Lundblad 1933a: 168-171, fig. 58; Brooks 1951: 416-418, figs. 60, 98; Lansbury 1965: 61. Material. - Sulawesi Tenggara: N8917, 4 ¢ 4 9 31 larvae; N8931, 3 6 5 Q. Pulau Buton: N8938, 16 & 21 Q. Distribution. - A very widespread species: E. India and Sri Lanka through China and Indonesia to Australia and Samoa. Anisops occipitalis Breddin Anisops occipitalis Breddin, 1905: 152. - Brooks 1951: 344-346, fig. 22; Lansbury 1965: 61-67, figs. 3-4, 7-8. Material. — Sulawesi Tenggara. N8908, 8 @ 15 9; N8917, 2 6; N8927, 4 & 7 9; N8931, 1 @ 1 9. Pulau Buton. N8938, 1 8 1 Q. Distribution. - Widespread through Indonesia to Northern Australia. Anisops paracrinata Brooks Anisops paracrinata Brooks 1951: 329-331, fig. 12. - Lansbury 1965: 57-58. > TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Material. - Sulawesi Tengah. SW of Luwuk, Totop camp along Batui River, 19-21 Oct 1989, Sul. 18, J. P. Duffels, 1 & (ZMA). Sulawesi Tenggara. Desa Kagunyala, N8908, 1 & 6 9; Jalan Asera, N8917, 26 & 23 9; Jalan Asera, N8918, 5 & 1 9; 20 km S Pomalaa, N8927, 4 &; 15 km S Pomalaa, N8931, 17 & 17 9. Pulau Buton. Desa Gareg-Gareng, N8938, 36 G © 2. Distribution. - Sumatera, Jawa, Bali, Sumbawa, Sulawesi, Buton, E. Australia. Anisops stali Kirkaldy Anisops stali Kirkaldy, 1904: 113, 132. - Brooks 1951: 319-322, figs. 9, 107; Lansbury 1965: 58. Material. - Sulawesi Tenggara. Desa Kagunyala, N8908, 2 9; 15 km S Pomalaa, N8931, 2 9. Remark. - Identification of these specimens un- certain since no males were available; they agree with specimens from Jawa. Distribution. - Widespread; the Philippines, Jawa, Sulawesi, Nusa Tenggara, Australia, Okin- awa. Anisops tahitiensis Lundblad Anisops tahitiensis Lundblad, 1934: 121-123, figs. 1-5. — Brooks 1951: 376-378, fig. 40; Lansbury 1964: 217- 218, fig. 7. Material. — Sulawesi Tenggara. 20 km E Kolaka, N8934, 4 6. Malaysia, Sabah. 20 km W of Sandakan, Sepilok-Laut. ML, 4 Nov. 1987, leg. J. Huisman & R. de Jong, 1 8 (RMNH). Remark. - The specimens were compared with a male from New Guinea and they differ only in the facial carina being slightly less prominent in the Sulawesi and Sabah specimens. Distribution. - Widespread: Andaman Islands, Borneo, Sulawesi, the Philippines, New Guinea, New Hebrides, Tahiti, Guadalcanal, Okinawa. Aphelonecta Lansbury This genus contains three species, one from Thailand and two from Borneo (Lansbury 1966). Aphelonecta alexis Lansbury Aphelonecta alexis Lansbury, 1965a: 328-330, figs. 2-18. - Lansbury 1966: 632. Material. - Malaysia, Sabah. 60 km W of Lahad Datu, Danum Valley, 14 Mar 1987 (B), leg. J. Huisman, 1 & (RMNH). Distribution. - Sabah and Sarawak. 58 Enithares Spinola A large, mostly Oriental genus. Thirty-three species are known from Malesia, of which seven occur in Sulawesi and one on Buton. There is an excellent revision of the Oriental species (Lansbury 1968). Key to the males of Enithares of Sulawesi and Buton 1. Embolium in ventral view (ventral ridge of hemielytron) greatly expanded anteriorly (fig. 21); pronotal humeral angles produced into broad knobs. Length about 9 mm (Sulawesi) El ahve seh ce i en Bai arr E. producta Lsb. - Embolium in ventral view not greatly ex- panded anteriorly; pronotal humeral angles nouproduced genant RN WAN eh een ae 2 Dinlensthiupito) in 1... eee eee een > — Length) Stmmyrorsmore m A 4 3. Head width twice the median length or less, anterior lobe of genital capsule bilobed due to a deep and broad median incision (widespread SPECIES) LENIHAN). ARE. E. bakeri Brks. — Head width just over twice its median length, anterior lobe of genital capsule not bilobed (P. Biak, possibly Sulawesi Tenggara) .............. O Sa VENE ROTTA E. vulgaris Lsb. 4. Length 12.5 mm, median length of head one- fourth of its length shorter than median length of pronotum, anterior margin of middle tibia straight to very slightly concave, posterior lobe of genital capsule broadly rodlike (Sulawesi Sélatan) wate. Po bassa: E. horvathi Kirk. - Length not over 12 mm; if apex of posterior lobe of genital capsule rodlike, then length not over 11 mm, anterior margin of middle tibia moreonlless convex sees Tee cee 5 5. Apex of posterior lobe of genital capsule rod- like (figs: 13516) G22 RE RARA RON 6 - Apex of posterior lobe of genital capsule rounded) (figsapl ZAIRE 7 6. Middle tibia not distinctly broadened (fig. 18), rod-like apices of posterior lobes of genital capsule rather short with small solid pegs on inner sides (fig. 13) (Sulawesi Tenggara) ...... AT Sio E. lansburyi sp. n. - Middle tibia distinctly broadened (fig. 20), rod- like apices of posterior lobes of genital capsule more elongate without pegs on inner sides (fig. 16) BMButon) Weeze E. skutalis sp. n. 7. Apical half of middle femora with very long hairs, covering ventral margin in inner view; genital capsule with a bilobed posterior lobe bearing an apicaudal tuft of setae (fig. 12) (Su- lawesi Tengah) ............. E. caesaries sp. n. - Middle femora without strikingly long hairs, posterior lobe of genital capsule not bilobed, withouttubtor setae (Migone OI PA on E. phenakismos sp. n. Enithares bakeri Brooks Enithares bakeri Brooks, 1948: 40, pl. 1, fig. 3. - Lansbury 1968: 384-385, figs. 91-95; Nieser & Chen, in press. Material. — Sulawesi Tenggara. Desa Kagunyala, N8908, 1 4 1 9; Aopa marsh, N8913, 1 9; Jalan Asera, N8917, 1 g; 15 km S Pomalaa, N8931, 4 9. Pulau Buton. Desa Gareg-Gareg, N8938, 1 & 1 9. Maluku. Bacan, Wayaua, eutrophic pool, HH382, 6-7 July 1985, leg. J. Huijbregts, 2 6 3 ® (RMNH). Distribution. - One of the few widespread spe- cies of Enithares in the area, occurring in the Phi- lippines, Sabah, Sulawesi, Maluku, Flores. Remarks. - The length of the females from Su- lawesi and Buton centers around 7.5 mm, which is slightly less, and of the specimens from the Maluku around 8.3, which is slightly more than indicated by Lansbury (1968). This suggests that the variability in size is greater than known up to now, and that there is significant variability between popula- tions. Enithares bakeri is smaller than most of its con- geners, and lives e.g. in village ponds together with the more common species of Anisops. Enithares hippokleides Kirkaldy Enithares hippokleides Kirkaldy, 1898: 73. — Lansbury 1968: 400-402, figs. 165-172 [redescription, synon- ymy |. Material. — Jawa, S. Muller [19th century], 1 6 2 9; G. Malang, Aug. 1935, 4 @ 2 © (all det. E. fruhstorferi by Brooks) (RMNH). Distribution. - Only known from Jawa. Enithares intricata Breddin Enithares intricata Breddin, 1905: 154. - Lundblad 1933a: 174-177, fig. 64, pl. 5, 21; Lansbury 1968: 404-405, figs. 185-190 [redescription, synonymy ]. Material. — N. Sumatra: Toba plateau, Tigadolok, 2°50°N 99°03’E, c. 150 m, 20 Jun 1972, leg. J. Krikken (21), 1 9 (RMNH). Distribution. - Sumatera and Jawa. Enithares producta Lansbury Enithares producta Lansbury, 1968: 368-369, figs. 15-21. NIESER & CHEN: Malesian Heteroptera Material. - Sulawesi Utara: Dumoga Bone N. P, sev- eral localities, incl. Tumpah river, Toraut river, Waterfall Creek, small creek near basecamp, leg. J. P. Duffels, J. Huijbregts, J. van Tol, total 13 & 14 9 (ZMA, RMNH). Sulawesi Tengah. 60 km SE Palu: Lore Lindu NP, Danau Tambing and brooklets, 1600 m, 7 Dec 1985, leg. J. van Tol, 1 ® (RMNH). Distribution. — Sulawesi. Enithares ripleyana Lansbury Enithares ripleyana Lansbury, 1968: 402-403, figs. 173- 178. Material. - Maluku. Bacan. Wayaua, logged forest, 5-16 July 1985, leg. J. Huijbregts, 1 @ (NC). Distribution. - Bacan, which is the present spell- ing of Batjan, the type locality of this species. Remarks. - This specimen agrees with the des- cription of E. ripleyana, except for its smaller size (7.3 mm), and the apical part of the posterior lobe of the genital capsule, which is inclined caudad. The dense tuft of setae on the genital capsule look solid when glued dry on a card. Enithares vulgaris Lansbury Enithares vulgaris Lansbury, 1968: 403-404. Material. — Sulawesi Tenggara. 20 km E Kolaka, N8934, 1 9. Distribution. - So far only known from the type series from P. Biak, situated east from the NW part of Irian Jaya. Remarks. - The size of the Sulawesi specimen is 9.1 mm, which indicates to E. vulgaris rather than to E. intricata Breddin. Males are needed for defin- itive identification. Enithares caesaries sp. n. (Figs. 12, 17) Type material. - Holotype @: Sulawesi Tengah. Luwuk area, Sungai Tikalalang, 20 Oct 1989, 89JvT26, leg. J. van Tol (RMNH). Description The description is based on the holotype only. Generally a rather large boat-shaped species, grea- test width at level of apex of scutellum. Measurements. — Length 10.3 mm, maximal width 4.03 mm, width of head 3.22 mm, humeral width of pronotum 3.81 mm, anterior width of vertex 1.10 mm, synthlipsis 0.70 mm. Colour. — Pale luteous, eyes dark brown, apical DO, TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 NN \ / (l Ue Figs. 12-16. Genital capsule in male Enithares. - 12, E. caesaries, holotype; 13, E. lansburyi, paratype; 14, E. hippokleides; 15, E. phenakismos, holotype; 16, E. skutalis, paratype. Scale bar 1 mm. 60 half of membrane smoky brown to blackish. Ven- tral side pale with rostrum, stripe along inner (cos- tal) margin of embolium, pilosity, spines, patches and stripes dark brown to blackish. Head and thorax. - Anterior margin of vertex in dorsal view truncate, hardly produced anterior to eyes. Greatest width of head 2.3 X its median length; median length of head and median length of pronotum subequal, somewhat longer than an- terior width of vertex. Humeral width of pronotum 3 X its median length, lateral margin slightly di- verging, posterior margin somewhat sinuate. Dor- sal margin of pronotal fovea slightly diverging behind eyes. Embolium only slightly expanded in anterior third. Nodal furrow about its own length from membranal suture. Legs. - Fore trochanter narrow posteriorly, with some long hairs, without nodule on ventral side, NIESER & CHEN: Malesian Heteroptera Figs. 17-20. Middle tibia of male Enithares. 17, E. cae- saries, holotype; 18, E. lans- buryi, paratype; 19, E. phe- nakismos, holotype; 20, E. skutalis, paratype. Scale bar 1 mm. meso-trochanter rounded. Mid-femoral hairs in apical half very long, covering ventral margin in inner view; mid tibia not broadened, slightly con- vex along anterior margin (fig. 17), outer claw of midleg normal. Ventral side. - Lateral edges of metaxiphus only slightly rounded and thickened, apical part with sinuate margins resulting in a sharply projecting apex. Connexiva of segments 1-3 with small black spines, not ridged. Male genitalia. - Genital capsule as in fig. 12; posterior lobe strongly sclerotized, its dorsal mar- gin incised, the distal top with a distinct tuft of hairs; anterior lobe less sclerotized, parameres very small, set in depressions in sides of capsule. Comparative notes In the key by Lansbury (1968) this species runs 61 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 1 a Fig. 21. Base of hemielytra and connexiva in Enithares, semidiagrammatical, in ventral view. Upper: greatly ex- panded anteriorly (as in E. producta), lower: not ex- panded. to E. alexis Lansb. in view of the midfemoral hairs, or, ignoring this character, to E. hippokleides Kirk. or E. timorensis Brooks. The last two species lack the midfemoral hairs, have also the middle tibia less hairy and the dorsal margin of the hindlobe of the genital capsule not incised. This species is also quite similar to E. lansburyi sp. n. (see the key). Etymology Caesaries (Latin: thick strands of hair), a noun in apposition, refers to the pilosity of the middle femur. Enithares lansburyi sp. n. (figs 13, 18) Type material. - Holotype &: Sulawesi Tenggara, small mountain stream 20 km E of Kolaka, N8934, 3 March 1989, leg. N. Nieser (ZMA). - Paratypes: 1 9 (ZMA); 3 & 3 Q (allotype) (NC, 1 & MBBJ). Description Shape. - Rather large boat-shaped species, grea- test width over pronotal humeral angles. Measurements. - Length @ 9.4-10.1 mm, @ 9.2- 10.0 mm; maximal width 3.7-3.9 mm; width of head 3.0-3.1 mm; humeral width of pronotum 3.7- 3.9 mm; anterior width of vertex 1.0-1.1 mm; syn- thlipsis 0.61-0.69 mm. Colour. - Pale form: Pale luteous, eyes dark brown, apical half of membrane and a large x- shaped spot at apex of scutellum smoky brown to blackish. Ventral side pale with proximal part of embolium, pilosity, spines, patches and stripes blackish. Dark form: anterior fourth of pronotum smoky brown, scutellum shiny black with, in fresh specimens, light greenish somewhat fluorescent stripes along lateral margins, in dried specimens these stripes become pale luteous. Hemielytra hya- line with apex of clavus, inner angle and outer band of corium and basal part of membrane black. Dor- sum of abdomen black, showing through hyaline 62 parts of hemielytra. Head and thorax. - Anterior margin of vertex in dorsal view truncate, slightly produced anterior to eyes. Greatest width of head 2.5 times its median length in male, just over two times its length in female; median length of head and median length of pronotum subequal, somewhat longer than an- terior width of vertex. Humeral width of pronotum just under three times its median length, lateral margins slightly diverging, posterior margin shal- lowly sinuate. Dorsal margin of pronotal fovea slightly diverging behind eyes. Embolium only slightly expanded in anterior third. Nodal furrow virtually straight with tip inclined cephalad, slightly less than its own length removed from membranal suture in male, distance and length subequal in female. Legs. - Fore trochanter nar- row posteriorly, with some long hairs, without nodule ventrally, middle trochanter rounded. Male middle tibia not broadened, slightly convex along anterior margin (fig. 18), outer claw of middle leg normal. Venter. - Metaxiphus with rounded and thick- ened lateral edges, apical part with sinuate margins resulting in a sharply projecting apex. Connexiva, segments 1 and 2 with small spines, not ridged. Male genitalia. - Genital capsule as in fig. 13, posterior lobe strongly sclerotized, long and stylus- like distally, the distal part with hairs and apically some spines, mainly located on the inner faces. Anterior lobe less sclerotized, parameres very small, set in depressions in sides of capsule. Comparative notes In the key by Lansbury (1968) this species runs to E. hippokleides Kirk., which lacks the elongate stylus-like apices of the hind lobes of the genital capsule. E. lansburyi is very similar to E. skutalis, but differs in the apex of the metaxiphus, which is more pronounced, and in the characters mentioned in the key. Etymology Named in honour of Dr. I. Lansbury for his excellent work on this genus and other Oriental Notonectidae. Biological notes E. lansburyi and E. skutalis were found in similar habitats, virtually stagnant, shaded pools asso- ciated with small streams in woodland. In each pool one or two specimens can be found, while larvae, especially the smaller ones, may be found in larger numbers and are of the same or of consecutive instars. We hypothize, that gravid Enithares fe- males seek out suitable ponds to deposit a number of eggs. The larvae start to use other kind of food, but in the end, when supplies become exhausted, resort to cannibalism. In this way the species stores the energy to produce a few adults temporarily in the larval population, which is also known in sev- eral species of Notonecta. Enithares phenakismos sp. n. (Figs. 15, 19) Material. - Holotype @: Sulawesi Tengah: Lore Lindu National Park, 10 km NE Gimpu, lower montane forest, Rano Rano, 1600 m, 15 Mar 1985, Sta. 42, J. P. Duffels & M. J. Duffels (ZMA). Description Only the holotype is known. Rather large boat- shaped species, greatest width probably at a level halfway scutellum (in the type right hemielytron somewhat broken and folded outward). Male. - Measurements. Length 11.9 mm, width of head 3.72 mm, humeral width of pronotum 4.3 mm, anterior width of vertex 1.40 mm, synthlipsis 0.98 mm. Colour. Pale luteous, eyes black, vertex light brownish, dark spot on abdominal dorsum at apices of clavi shining through hemielytra, which are opaque except for apical, hyaline half of mem- brane. Ventrally pale with stripes, pilosity and spines mostly dark brown to blackish. Head and thorax. Anterior margin of vertex in dorsal view rounded, hardly produced anterior of eyes. Greatest width of head twice its median length; median length of head and median length of pronotum subequal, only slightly longer than anterior width of vertex. Humeral width of pro- notum three times its median length, lateral mar- gins slightly diverging, posterior margin nearly straight in median part, gently curved anteriorly in lateral parts. Dorsal margin of pronotal fovea con- verging behind eyes. Embolium only slightly ex- panded in anterior third. Nodal furrow about its own length removed from membranal suture. Legs. Fore trochanter posteriorly narrow, with some long hairs, without nodule ventrally, meso- trochanter rounded. Mid-femoral hairs normal, with a row of short black hairs near suture with trochanter, but lacking the small patch of short black bristles at its apex, which are visible in most specimens of E. hippokleides, middle tibia not broadened, slightly convex along anterior margin (fig. 19), outer claw of middle leg normal. Venter. Sides of metaxiphus somewhat rounded and thicken, apex long and sharply projecting. Connexiva of segments 1-3 with small black spines, not ridged. Genital capsule as in fig. 15, posterior lobe with sinuate posterior margin. NIESER & CHEN: Malesian Heteroptera Comparative notes This species runs to E. hippokleides Kirk. in the key by Lansbury (1968). At first sight it looks like an oversized specimen of E. hippokleides, includ- ing a similarity of the genital capsule and the pilo- sity of the legs. On closer inspection there are several small structural differences, viz. head width / synthlipsis about 6 in E. hippokleides and less than 4 in E. phenakismos, anterior width vertex / synthlipsis about 2 in E. hippokleides and 1.5 in E. phenakismos, dorsal margin of pronotal fovea slightly diverging posteriorly in E. hippokleides and converging in E. phenakismos, apex of meta- xiphus longer in E. phenakismos than in E. hippo- kleides. Characters to distinguish E. phenakismos from other new species described in this paper can be found in the key. E. timorensis Brooks, which is considered to be closely related to E. hippokleides, is even smaller (length of male up to 9.5 mm) and has the posterior lobe of the genital capsule not sinuate. Etymology Phenakismos (Gr. deceit), a noun in apposition, refers to the close similarity to E. hippokleides Kirk. Enithares skutalis sp. n. (figs. 16, 20) Type material. - Holotype &: Pulau Buton, 16 km E Baubau, small stream in wet forest, 10.III.1989, N8944, leg. N. Nieser (ZMA). - Paratypes: 1 @ with same data as holotype (NC); Buton, small mountain stream, N8941, 2 8 (NC), 1& (MBBJ). Description Male. - Shape. Rather large, boat-shaped spe- cies, greatest width across hemielytra just behind pronotal humeral angles. , Measurements. Length 10.4-10.8 mm, maximal width 4.10-4.15 mm, width of head 3.25 mm, hu- meral width of pronotum 4.00-4.05 mm, anterior width of vertex 1.15-1.20 mm, synthlipsis 0.65- 0.68 mm. Colour. Pale luteous, eyes dark brown, apical half of membrane smoky brown, spot just caudally of scutellum light smoky brown. Ventral side pale with pilosity, spines, patches and stripes blackish. Head and thorax. Anterior margin of head in dorsal view nearly straight. Greatest width of head 2.5 X its median length; median length of head, median length of pronotum and anterior width of vertex subequal. Humeral width of pronotum more than 3 X its median length. Pronotum with lateral margins diverging, posterior margin straight; dor- 63 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 sal margin of pronotal fovea directed virtually straight caudad behind eyes. Embolium only slightly expanded in anterior third. Nodal furrow virtually straight with tip inclined cephalad, slightly less than its own length removed from membranal suture. Legs. Fore trochanter narrow posteriorly, co- vered with thick long hairs, without nodule ven- trally, mesotrochanter rounded. Male middle tibia slightly broadened, convex along anterior margin (fig. 20), outer claw of middle leg normal. Venter. Metaxiphus with distinct rounded and thickened lateral edges, apical part triangular with virtually straight lateral margins. Connexiva of segments 1 and 2 with small black spines, not ridged. Genital capsule as in fig. 16, posterior lobe strongly sclerotized, long, hairy and distally stylus- like. Anterior lobe less sclerotized, parameres very small, set in depressions in sides of capsule. Comparative notes This species runs to E. hebridensis Lansb. or E. hippokleides Kirk. in the key by Lansbury (1968), which, however, both lack the elongate stylus-like apex of the hind lobe of the genital capsule, and differ in the shape of the male middle tibia. The genital capsule of E. skutalis has a structure similar to E. horvathi and E. lansburyi, but in detail these are quite different. Besides, E. horvathi is two mm longer and has the anterior margin of the male middle tibia slightly concave (convex in E. skuta- lis). Etymology Skutalis (Gr. baton), a noun in apposition, refers to the rodlike shape of the apex of the posterior lobe of the genital capsule. Nychia Stàl Genus with only a few, mostly poorly known species. Apparently only one species in Malesia (Lansbury 1985). Nychia sappho Kirkaldy Nychia marshalli var. sappho Kirkaldy, 1901a: 809-810. Nychia malayana Lundblad, 1933a: 148-155, figs. 49-51. Nychia sappho; Lansbury 1985: 4-8, figs. 7-21. Material. - Sulawesi Tenggara: Wawanggole, N8901, 2 8 4 9; Sungai Sampara, N8909, 3 & 3 9; 2 km E Sungai Sampara, N8910, 1 9; Aopa marsh, N8913, 7 6 3 9; Jalan Asera, N8916, 1 9. Pulau Buton: Road to Lawele, N8939, 5 & 5 9, all brachypterous. Distribution. - Malaysia, Indonesia, New Guinea and Northern Australia. 64 REFERENCES Breddin, G., 1905. Rhynchota Heteroptera aus Java. - Mitteilungen aus dem Naturhistorischen Museum in Hamburg 22: 111-159. Brooks, G. T., 1948. New species of Enithares (Hemip- tera, Notonectidae). - Journal of the Kansas Entomo- logical Society 21: 37-54, 3 plates. Brooks, G. T., 1951. A revision of the genus Anisops (Notonectidae, Hemiptera). - Science Bulletin, Uni- versity of Kansas 34: 301-519. Fieber, F. X., 1851. Rhynchotographien III. Die Gruppe der Notonectae. - Abhandlungen der königl. böh- mischer Gesellschaft der Wissenschaften 5 (7): 469- 486. Hale, H. M., 1924. Studies in Australian aquatic Hemip- tera III. Nepidae. - Records of the South Australian Museum 2: 503-520, pl. 24-26. Kirkaldy, G. W., 1898. Deux especes nouvelles du genre Enithares Spin. et notes hemipterologiques diverses. - Revue d’Entomologie Caen 17: 72-73. Kirkaldy, G. W., 1901. Miscellanea Rhynchotalia. - Ento- mologist 34: 5-6. Kirkaldy, G. W., 1901a. On some Rhynchota, principally from New Guinea. (Amphibicorisae and Notonecti- dae). - Annali del Museo Civico do Storia Naturale G. Doria 20: 804-810. Kirkaldy, G. W., 1904. Uber Notonectiden. - Wiener Entomologische Zeitung 23: 93-135. Lansbury, I., 1964. Some observations on the Notonecti- dae (Hemiptera-Heteroptera) of Viet-Nam and adja- cent regions. — Annales Zoologici (Warszawa) 22: 203-219. Lansbury, I., 1965. Notes on the species of the genus Anisops Spin. (Hemiptera - Heteroptera) of Java. - Annales Zoologici (Warszawa) 23: 57-68. Lansbury, I., 1965a. A new tribe and genus of Notonec- tidae (Heteroptera: Notonectidae) from Borneo. - Pacific Insects 7: 327-332. Lansbury, I., 1966. Notes on the genus Aphelonecta (He- miptera - Heteroptera: Notonectidae). - Pacific In- sects 8: 629-632. Lansbury, I., 1967. Comparative morphology of the male Australian Nepidae (Hemiptera: Heteroptera). - Australian Journal of Zoology 15: 641-649. Lansbury, I., 1968. The Enithares (Hemiptera - Heterop- tera: Notonectidae) of the Oriental region. — Pacific Insects 10: 353-442. Lansbury, I., 1972. A review of the Oriental species of Ranatra Fabricius (Hemiptera - Heteroptera: Nepi- dae). - Transactions of the Royal Entomological So- ciety of London 124: 287-341. Lansbury, I., 1973. A review of the genus Cercotmetus Amyot & Serville, 1843 (Hemiptera - Heteroptera: Nepidae) - Tijdschrift voor Entomologie 116: 83-106. Lansbury, I., 1978. Comments on the species concept in some Australian Anisops Spinola (Hemiptera: Noto- nectidae). - Memoirs of the National Museum of Victoria 39: 101-115. Lansbury, I., 1985. Notes on the identity of Nychia Stal (Hemiptera - Heteroptera: Notonectidae) in Austra- lia. - Beagle 2: 1-9. La Rivers, I., 1970. Two new species of Laccocoris from British North Borneo (Hemiptera: Naucoridae: Lac- cocorinae). - Proceedings of the Entomological So- ciety of Washington 72: 496-499. La Rivers, I., 1971. Catalogue of taxa described in the family Naucoridae (Hemiptera). - Biological Society of Nevada Memoir 2: 65-99. Leong, C. Y., 1962. The life-history of Anisops breddini Kirkaldy (Hemiptera, Notonectidae). - Annals and Magazine of Natural History (Series 13) 5: 377-383. Lundblad, O., 1933. Some new or little known Rhynchota from China. - Annals and Magazine of Natural His- tory (Series 10) 12: 449-464, pl. 13. Lundblad, O., 1933a. Zur Kenntnis der aquatilen und semiaquatilen Hemipteren von Sumatra, Java und Bali. - Archiv fiir Hydrobiologie, Supplement 12: 1- 195, 263-489, pls. 1-21, figs. 1-142. Lundblad, O., 1934. Aquatic and semiaquatic Heteroptera of Tahiti. - Bulletin of the B. P. Bishop Museum 113: 121-126. Montandon, A. L., 1897. Hemiptera Cryptocerata. Fam. Naucoridae. Sous-fam. Laccocorinae. - Verhandlun- gen der K.-K. zoologisch-botanischen Vereins Wien 47: 435-454. Montandon, A. L., 1911. Nouvelles espèces d’Hydroco- rises appartenant aux collections du Muséum Ento- mologique de Berlin. - Bulletinul Societatii de Stiinte 20: 83-93. Nieser, N. & P. Chen, in press. Notes on Indonesian waterbugs (Nepomorpha & Gerromorpha). - Storkia Polhemus, D. A., 1986. A review of the genus Coptocatus Montandon (Hemiptera: Naucoridae). - Pan Pacific Entomologist 62: 248-256. Polhemus, D. A. & J. T. Polhemus, 1988. The Aphelochei- rinae of tropical Asia (Heteroptera: Naucoridae). — Raffles Bulletin of Zoology 36: 167-300. Stal, C., 1854. Nya Hemiptera. - Ofversikt af Kongliga Svenska Vetenskaps-Akademiens Förhandlingar 11: 231-255. Stal, C., 1859. Hemiptera. - Kongliga Svenska Fregatten Eugenies Resa Omkring Norden, Zoologi 4: 219-298. Stal, C., 1861. Nova methodus familias quasdam Hemip- tera disponendi. - Ofversikt af Kongliga Svenska Ve- tenskaps-Akademiens Förhandlingar 18: 195-212. Stal, C., 1870. Hemiptera insularum Philippinarum. - Ofversikt af Kongliga Svenska Vetenskaps-Akade- miens Förhandlingar 27: 607-776, pls. 7-9. Van Tol, J. et al, in prep. Report of the 1989 RMNH Expedition to Sulawesi. Whitten, A. J., M. Mustafa & G. S. Henderson, 1988. The ecology of Sulawesi. - Gadjah Mada University Press, Yogyakarta: 1-777, figs., plates. Received: 12 October 1990 Revised version accepted: 30 March 1991 NIESER & CHEN: Malesian Heteroptera APPENDIX 1 Localities sampled by N. Nieser, Indonesia, Su- lawesi Tenggara, 1989. N8901-N8920: Kabupaten Kendari N8901-N8902: Kecamatan Wawotobi N8901. Wawonggole, Sungai Anggoro, 20 Feb 1989. Quietly flowing stream in open woodland, width 4-5 m, depth up to 1 m, dark brown water; vegetation hanging from the banks in the water. N8902. Wawongole. 20 Feb. 1989. Pond with abundant Nymphaea in marshy fields; shore vegetation hanging into the water. N8903-N8912: Kecamatan Kendari N8903. Eastern road to Lalimboee, 3 km outside Kendari, 21 Feb. 1989. Small stream, width 2-3 m, current 15m/m, slower in ponded sites, bottom sandy, some detergents, but clear. N8903A. Same site, puddle at edge. N8904. Eastern road to Lalimboe, 4 km outside Kendari, 21 Feb. 1989. Turbid, near stagnant stream, width 1.5 m, depth up to 1 m. N8905. Desa Kagunyala, 21 Feb. 1989. Trench. N8906. Desa Kagunyala, 21 Feb. 1989. Pond covered with Azolla and Lemna. 100 X 10 m, depth 0.05 m, bottom clay and mud. N8907. Desa Kagunyala, 21 Feb. 1989. Pond with some Lemna, 15 X 5 m, depth 0.05 m, bottom clay and mud. N8908. Road outside Desa Kagunyala, 21 Feb. 1989. Pond, 100 X 50 m, turbid, beige, grass-like vegetation along one bank. Many Cybister and Dytiscus. N8909. Sungai Sampara, second bridge along road Ken- dari-Wawotobi, 22 Feb. 1989. River, sampled along bank at bridge, water turbid, beige, sand bottom. N8910. About 2 km E of N8909, 22 Feb. 1989. Puddle in dry stream bed in savannah-like area at foot of hill with monsoon forest. N8911. Small stream about 8 km E of N8909, 22 Feb. 1989. Width 3 m, depth 0.6 m, water turbid, beige, bottom sand and loam. Current up to 5 m/min. N8912. Puddle high on banks of N8911, 22 Feb. 1989. Dimensions 10 X 3 m, depth 0.4 m. N8913-N8915: Kecamatan Lembuya N8913. Aopa marsh, 23 Feb. 1989. Bay in marsh at office building, grass-like vegetation on banks, Ceratophyl- lum in water, water light brown but clear. N8914. Road Lembuya-Palangga, desa Lamooso, 23 Feb. 1989. Sungai Simbangi. Open loamy and sand bottom. Slow current. N8915. Road Lembuya-Palangga, 23 Feb. 1989. Swampy banks of stream in savannah. N8916-N8920. Kecamatan Asera. N8916. Jalan Asera, first stream drom bridge over S. Sampara, 24 Feb. 1989. Riffles with current 15-20 m/min. Rhagovelia. N8917. Pool in meadow near N8916, 24 Feb. 1989. Di- mensions 4 X 2.5 m, depth up to 0.3 m, water turbid, beige, grass-like vegetation and Polygonum. N8918. Flooded banks of second stream, 24 Feb. 1989. Meadows mainly with Cyperaceae, water turbid, beige. N8919. Same road, but here named Jalan Paku Jaya, fifth stream at slightly higher elevation, 24 Feb. 1989. At bridge, slow current, bottom sand and clay. N8920. Same road, fourth stream, coming from wood- 65 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 land, 24 Feb. 1989. Ptilomera in shade of trees, Lim- nogonus in exposed puddle aside of current. N8921-N8934: Kabupaten Kolaka N8921. Kecamatan Kolaka N8921. Sungai Kolaka, 27 Feb. 1989. Upstream of Kol- aka, river, bottom sand and pebbles, current variable. Ochterus and Hydrometra on mudflat. N8921 A. Same site, places with current. N8921B. Same site, ponded sites. N8922-N8924: Kecamatan Mowewe. N8922. Sungai Mowewe, 28 Feb. 1989. Lowland stream, 2.5-10 m wide, in woodland with fields and settle- ments closeby. Water clear, colourless. Bottom sample at shallow, partly shaded places with moderate cur- rent, bottom coarse sand and small pebbles. N8922A. Same site, pothole with undercut bank, tree roots, floating plant debris, depth over 1 m. N8923. Small stream in fields and gardens near N8922, 28 Feb. 1989. Width 0.5-1 m, depth 0.1-0.3 m, bottom mostly pebbles, rather fast current. N8924. Puddles on path at N8922-N8923, 28 Feb. 1989. N8925-N8931: Kecamatan Kolaka. N8925. Tamborasi, 1 Mar 1989. Subterraneous stream flowing into small creek. N8926. Between Tamborasi and Wolo, 1 Mar. 1989. Mountain stream through cocoa plantation. Ptilomera and Rhagovelia on more quiet sites behind boulders. Limnogonus on puddle on banks. N8927. About 20 km S of Pomalaa, 2 Mar. 1989. Pool at edge of marsh, 2 X 2 m, depth up to 0.7 m, bottom loam, water turbid, beige. Juncus, Chara, herbaceous plants. N8928. Small stream in same marsh as N8927, 2 Mar. 1989. Water clear, light brown, flowing through dense marsh vegetation, sample at culvert under road, width 1 m, current c. 10 m/min, depth up to 0.7 m, bottom sand with pebbles. N8929. Pond at edge of marshy forest. 2 Mar. 1989. Nymphaea, Juncus. N8930. Small stream in marsh, 2 Mar. 1989. Open part with flooded banks. Grass-like vegetation along banks with considerable amount of epiphytic algae. Most species from flooded banks, but Rhagovelia from the stream. N8931. C. 15 km S of Pomalaa, 2 Mar. 1989. Sand pits at edge of marsh. Water clear to somewhat turbid, light brown, depth up to 0.5 m. N8932-N8933: Kecamatan Mowewe. N8932. Road to Mowewe, 3 Mar. 1989. Narrow stream at edge of marsh, water dark brown, bottom with much muddy detritus and plant debris, some Azolla. N8933. Wide stream near N8932, 3 Mar. 1989. Sample from shallow bay without current, soft loamy bottom, much debris from palmtree leaves. N8934. Kecamatan Kolaka. N8934. Road to Kendari, 20 km E of Kolaka, 3 Mar. 1989. Small mountain stream in woodland, 1-1.5 m wide, depth very variable, potholes up to 1 m, current very variable, small waterfalls and ponded sites. N8935-N8944. Kabupaten Buton N8935. First stream crossing road from Baubau to the north, 8 Mar. 1989. Downstream of man-made water- fall, width 5 m, depth and current velocity variable, water colourless, somewhat turbid. 66 N8936. Desa Gareng-gareng, 8 Mar. 1989. Pool with many small fish. N8937. Desa Gareng-gareng, 8 Mar. 1989. Pool with very few fish. N8938. Desa Gareng-gareng, 8 Mar. 1989. Dirty pond without fish. N8939. Road to Lawele, 9 Mar. 1989. Mangrove swamp E of road, bottom with thin layer of mud and man- grove roots, marshy vegetation. N8940. Lower course of small stream, 9 Mar. 1989. Bot- tom bare grey sand and mud, some plant debris, slug- gish. N8941. Small mountain stream close to the sea, 9 Mar. 1989. Bottom rocky without sand or pebbles, vary from small waterfalls to nearly stagnant pools, no water vegetation, but banks with dense cover of mosses. N8942. Road to Parjowiro, about 15 km E of Baubau. 10 Mar. 1989. Small cascade-stream, water turbid, and somewhat milky colour. N8943. Pool at edge of N8942, 10 Mar. 1989. N8944. Road to Parjowiro, about 16 km E of Baubau, 10 Mar. 1989. Small stream, anastomosing, ponded sites, marshy forest, many buttressed trees, hilly country. N8945-N8949. Kabupaten Kendari. N8945-N8949. Kecamatan Kendari. N8945. Teluk Kendari, 13 Mar. 1989. Small stream flow- ing through coconut plantation into bay, bottom sand and clay. N8946. Teluk Kendari, 13 Mar. 1989. Well near coconut plantation. N8947. Teluk Kendari, Pulau Bungku Toko, 13 Mar. 1989. Puddles in mangrove. N8948. Teluk Kendari, close to sea. 13 Mar. 1989. Pool with seawater behind sand wall overgrown with Ipomoea. N8949. Teluk Kendari. 13 Mar. 1989. Well protected by masonry, near N8948. APPENDIX 2 Localities on Sulawesi and Borneo from recent col- lections made by J. P. Duffels, J. Huisman, R. de Jong and J. van Tol. Borneo 14 Mar 1987 - Sabah. 60 km W of Lahad Datu. Danum Valley near Danum Valley Field Centre, brooklet at West 3. Sample B. Alt. 220 m asl. 4°58’N 117°48’E. (J. Huisman). 23 Mar 1987 - Sabah. 60 km W of Lahad Datu. Danum Valley near Danum Valley Field Centre, streamlet near West 11 / North 5. Sample A. Alt. 220 m asl. 4°48’N 117°48E. (J. Huisman). 3 Apr 1987 - S Sabah. 105 km S of Beaufort: Long Pasia area. Sungai Maga near confluence Sg. Pasia. Alt. 1210 m asl. Larger fast running stream in untouched lower montane evergreen rain forest. Large boulders, rapids. 4°26'N 115°40’E. (J. van Tol). 8-9 Apr 1987 -S Sabah. 105 km S of Beaufort: Long Pasia area. Sungai Ritan. Alt. 1160 m asl. Undisturbed ever- green tropical rain forest. 4°24'N 115°42'E. (J. van Tol & J. Huisman). 4 Nov 1987 - Sabah. 20 km W of Sandakan. Sepilok Laut near resthouse. ML light. 5°49’N 118°0G'E. (J. Huis- man & R. de Jong). 23 Nov 1987 - Sabah. 16 km NE Tenom: Agricultural Research Station. Sungai Segalan. At light. 5° 12° N 115°59'E. (J. Huisman & R. de Jong). Sulawesi 23 Feb 1985 — Sulawesi Utara. Dumoga-Bone NP. Project Wallace. Edward's Camp. Tumpah river. 900 m asl [recte 600 m]. (J. P. Duffels). 15 Mar 1985 - Sulawesi Tengah. Lore Lindu National Park. 10 km NE of Gimpu. Lower montane forest. Rano Rano. Alt. 1600 m asl. Duffels Sta. 42 (J. P. & M. J. Duffels). 23 Apr 1985 - Sulawesi Utara. Dumoga-Bone NP. Project Wallace. Waterfall Creek, tributary of Tumpah river. Sample B. c. 225 m asl. UTM WL9768 [ recte XL0064]. 0°35’N 123° 54’E. (J. van Tol). 23 May 1985 — Sulawesi Utara. Dumoga-Bone NP. Pro- ject Wallace. Tumpah river near confluence Toraut. Sample A. c. 210 m asl. UTM XL0063. 0°34’N 123°54E (J. van Tol). 3 Jun 1985 - Sulawesi Utara. Dumoga-Bone NP. Project NIESER & CHEN: Malesian Heteroptera Wallace. Tumpah river near Edward's subcamp. Sam- ple A. Alt. c. 600 m asl. UTM WL9365. 0°35’N 123°51E. (J. van Tol). 4 Jun 1985 - Sulawesi Utara. Dumoga-Bone NP. Project Wallace. Brooklet c. 2 km west of Edward’s subcamp. Sample A. c. 700 m asl. UTM WL9269. (J. van Tol). 6 Dec 1985 - Sulawesi Tengah. 50 km SE of Palu: Lore Lindu National Park. Sopu valley near Dongi Dongi. Sample B. Alt. 950 m asl. Rainwater puddle without vegetation, near logging road. UTM SJ86. 1°13'S 120° 11'E (J. van Tol & J. Krikken). 7 Dec 1985 - Sulawesi Tengah. 60 km SE of Palu: Lore Lindu National Park. Danau Tambing and brooklets. Sample A. Alt. 1600 m asl. Pandanus. 1°20'S 120° 15’E. (J. van Tol). 8 Dec 1985 — Sulawesi Tengah. 50 km SE of Palu: Lore Lindu National Park. Sopu river near Dongi Dongi. Sample A. Alt. 950 m asl. UTM=SJ86. 1°13'S 120° 11’E (J. van Tol). 19-21 Oct 1989. - Sulawesi Tengah. SW of Luwuk: Totop camp along Batui river. Alt. 120 m asl. 1°09'S 122°31’30”E. Lowland rainforest. Sample Sul. 18 (J. P. Duffels). 20 Oct 1989. — Sulawesi Tengah. Luwuk area. Sungai Tikalalang. Sample 89JvT026. (J. van Tol). 67 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 BOOK REVIEW Medvedev, G. S. (editor-in-chief), 1990. Keys to the In- sects of the European Part of the USSR. Volume IV. Lepidoptera, Part 2. - Keys to the fauna of the USSR, 130. English Translation [originally published in Rus- sian, Leningrad, 1981]. - E. J. Brill, Leiden. 1092 pp, 747 figs. [ISBN 90 04 08926 8, price 280 Dutch guilders, ca US $ 140]. This book is the second volume on Lepidoptera translated into english. The first was published in 1988 and reviewed by the late A. Diakonoff in this journal (vol. 132: 162). While the first volume dealt with the primitive Lepidoptera, the Tortrici- dae and some primitive Macrolepidoptera, this vo- lume treats most of the remaining so-called mi- crolepidopteran families, covering the superfam- ilies Tineoidea (incl. Gracillariidae and related families, but excluding the Psychidae, which were treated in vol. 1), Yponomeutoidea and Gelechioi- dea (excluding the Coleophoridae). In total about 1280 species from the European part of the USSR and adjacent countries are covered. Keys are pro- vided up to species level and genitalia of many species are illustrated. Also some adults are illus- trated in line-figures. Although this is a review of the translation, it also presents an opportunity, being unable to read russian, to review the original contents, bearing in mind that it represents the knowledge of about 1980. Preparing identification keys for a large order as the Lepidoptera for the huge territory of the Euro- pean part of the USSR is in itself a large project, which is unprecedented outside the USSR. The keys heavily lean on previously published revisions and keys, mostly published outside the USSR. However, gathering such information is worth- wile, especially for the non-specialist, where the originals are so scattered over journals and books. For some families there is also original input, in particular figures of genitalia of species never illus- trated before. Regrettably, many male genitalia have been illustrated in lateral view, where it is now general practice to illustrate male genitalia in taxonomic papers in ventral view, with the valvae spread out. This will make comparison of standard genitalia preparations with the figures difficult, and could lead to misidentifications. 68 The quality of the various families differs greatly, some are clearly treated by specialists, and contain original information, others are in fact almost copies of foreign literature, like the family Elachistidae, which is merely an abstract of Volume 6 of Fauna Entomologica Scandinavica by Trau- gott-Olsen & Nielsen (now sold by the same pub- lisher as this translation!). Important original treatments are the Gracillariidae by Kuznetsov, the Momphidae and related families by Zagulyaev and Sinev and the Oecophoridae by L'vovskii. The chapter on Gelechiidae by Piskunov is important, because it is the only currently available treatment of European Gelechiidae. Unfortunately, the qual- ity is below expectations, since many species are excluded from the keys, the genitalia illustrations are a collection of copies from other sources, in- cluding some very poor ones, and complex original figures in lateral view. Although it is understandable from the point of costs, what I feel is most missing are colour plates of adult moths. Identification of moths should start with comparison of colour patterns, which is so much easier than to start making preparations of wing venation and genitalia. With this book, iden- tification of an unknown genus is often almost impossible without referring to the genitalia. I am not able to comment on the quality of the translation, but would like to make two general remarks. Firstly, the reproduction of the figures is poor. Despite the poor quality of paper, the figures in the Russian text are far superior. Secondly, it is to be regretted that the translators have chosen to translate the russian common host-plant names into common english names. Although there is a list of these names and scientific names at the end of the book, it would have been much easier to include scientific names, where many users will not be familiar with english common plant names. In conclusion, this translation is an important review of European Microlepidoptera, which tem- porarily will fill the gap between the older plate books like ‘Spuler’ and the modern treatments of ‘Microlepidoptera Palaearctica’, but it should be used which much care, since it is definitely not complete and contains some erroneous informa- tion. Those, who use the book merely for the illus- trations of genitalia, are beter off with the original russian edition, which is also much cheaper. [E. J. van Nieukerken] R. PUPLESIS!, S. SEKSJAEVA? & V. SRUOGA! 'Zoologyos katedra VPI, Vilnius ‘Zoological institute, Academy of Sciences, Leningrad LEAF-MINING LEPIDOPTERA (NEPTICULIDAE, BUCCULATRICIDAE, GRACILLARIIDAE) FROM ULMUS IN NORTHERN CASPIYA (KASPIA) Puplesis, R., Seksjaeva, S. & V. Sruoga, 1991. Leaf-mining Lepidoptera (Nepticulidae, Bucculatricidae, Gracillariidae) from Ulmus in northern Caspiya (Kaspia). - Tijdschrift voor Entomologie 134: 69-73, figs. 1-9. [ISSN 0040-7496]. Published 1 July 1991. Three leaf-mining species from elm (Ulmus carpinifolia Rupp. ex Suckow): Stigmella kazakhstanica Puplesis sp. n. (Nepticulidae), Bucculatrix caspica Puplesis & Sruoga sp. n. (Bucculatricidae) and Phyllonorycter schreberella (F.) (Gracillariidae) are recorded from the northern Caspiyan region (USSR). The new species are described, genitalia and mines are illustrated. Phyllonorycter schreberella (F.) is diagnosed and its male genitalia are figured. Correspondence: Dr. R. Puplesis, Zoologijos katedra VPI, g. Studentu 39, Vilnius 34, Lithuania. Keywords. - Nepticulidae; Bucculatricidae; Gracillariidae; leaf-mining Lepidoptera; northern Caspiya. The large northern Caspiyan (=Kaspian) region is generally characterized by a strong continental climate and by arid landscapes. Except oases by the rare rivers and settlements, there are only dry steppes, semideserts and deserts. The abundance of salt lakes, saline soils (solanchaks and solonetzes) is very characteristic. The vegetation of most of these biotopes is extremely poor, usually without woody plants. Grass cover is thin, the vegetation consists mainly of different worms-woods (Artemisia spp.) and cereals, with worm-woods dominating. The sum of active temperature equals 2800-3400 °C with a wetting coefficient of 0.15- 0.10 (Tushinskij & Davydova 1976). In oases near the settlements the elm (U/mus carpinifolia Rupp. ex Suckow) is the dominating tree. In a few cases it is the only tree present. Hardly any published data on leaf-mining Lepi- doptera were available from these regions. Only one Stigmella (Nepticulidae) specimen deposited in the Zoological institute (Leningrad), reared by G. Lindeman from Ulmus sp., from Dzhanibek (Western Kazakhstan), was known. It belongs to a new species. During investigations in early August 1988 in the localities (fig. 1) Baskunschak (Astrakhan Re- gion), Gurjew (Gurjew Region), Beyneu (Man- gyschlak Region), Kara-Kalpakiya and Kungrad (Kara-Kalpakskaya ASSR), mines of three leaf- mining Lepidoptera were discovered. Bucculatrix caspica Puplesis & Sruoga sp. n. was found in the largest numbers. Numerous cocoons and empty mines were collected from Ulmus carpinifolia in all investigated localities, Gurjew and Kungrad. Our later study of 1988 in neighboring Turkmeniya showed its absence from the Central Asiatic part of the USSR. Abundant mines of Phyllonorycter schreberella (F.), containing larvae or pupae were collected on Ulmus carpinifolia in Baskuntschak. In other places of the Caspiyan region, this species remains yet unrecorded. One empty mine of Nep- ticulidae was found on Ulmus carpinifolia in Bas- kuntschak, i. e. about 140 km South of Dzhanibek, where the above mentioned Stigmella specimen on Ulmus sp. was collected by Lindeman in 1966. It is therefore considered to belong to the same species. Ulmus spp. (including U. carpinifolia Rupp. ex Suckow) are known to be host plants for Gracilla- riidae (Kuznetzov 1981), Bucculatricidae (Seks- jaeva 1981) and Nepticulidae (Johansson & Nielsen 1990). No other lepidopterous miners on other plants have been found in this region. The type specimens are deposited in the collec- tion of the Minological research laboratory at the Department of Zoology of the Pedagogical insti- 69 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Fig. 1. Distribution map of leaf-mining Lepidoptera in northern Caspiya: Stigmella kazakhstanica (rectangles), Bucculatrix caspica (dots), Phyllonorycter schreberella (triangle). tute, Vilnius, Lithuania (MRL) and in the collection of the Zoological institute of the USSR Academy of Sciences in Leningrad (ZIAS). TAXONOMIC PART Nepticulidae Stigmella kazakhstanica Puplesis sp. n. (figs. 1-3) Type material. - Holotype &: USSR, western Kazakh- stan (Kazakh SSR), Dzhanibek, larva on leaf of Ulmus sp. (probably U. carpinifolia), vi.1966, leg. G. Lindeman (ZIAS). Leaf-mine (no type material): USSR, Astrakhan Region, Baskuntschak, on Ulmus carpinifolia, fresh mine, 4.viii.1988, leg. R. Puplesis et V. Sruoga (MRL). Diagnosis. - Belongs to the Stigmella ulmivora group. It is very similar to S. ulmiphaga (Preis- secker) and S. ulmivora (Fologne). In the male genitalia it is easily distinguished by the valvae, which are abruptly broadened at the base. Cornuti not numerous, lateral lobes of vinculum broad in contrast to S. ulmivora (Johansson & Nielsen 1990). Description Male. - Frontal tuft pale orange. Antennae pale brown. Eye-caps and collar cream. Palpi cream. Thorax and forewings uniform, greyish brown. Ci- lia and hindwings approximately as forewings in colour. 70 Fig. 2. Male genitalia of Stigmella kazakhstanica, holo- type (scale 0.1 mm). Fig. 3. Mine of Stigmella kazakhstanica on Ulmus sp., from which the holotype was reared. Female unknown. Male genitalia (fig. 2). - Valva narrow medially and apically, strongly broadened basally. Transtilla with short sublateral processes. Juxta present, its form resembles a triangle apically. Uncus approx- imately with trapezium shape, with slightly taper- ing angles. Gnathos with two thin and long poste- rior processus. Transverse bar of gnathos narrow. Aedeagus narrowing in middle (in lateral view). Vesica with two groups of cornuti; one of which is larger and situated in middle, a smaller group is si- tuated apically. Cornuti more or less in form of scle- rotized spines, some cornuti have different forms. Biology. — Larvae in June and August, mining leaves of Ulmus carpinifolia Rupp. ex Suckow. Mine a contorted gallery (fig. 3). Early mine com- pletely filled with brownish frass; later gradually widening with frass coiled, leaving wide clear mar- gins; finally frass in a dark narrow line. Distribution (fig. 1). - Only known from the lower Volga valley. Bucculatricidae Bucculatrix caspica Puplesis & Sruoga sp. n. (figs. 1, 4-8) Type material. - Holotype &: USSR, Astrakhan Re- gion, Baskuntschak, N 4170, cocoon on leaf Ulmus car- pinifolia, 4.viii.1988, ex p. 7.viii.1988. Paratypes: 1 6, 1 Q, same data as holotype, ex p. 7-15.viii.1988; 1 8, same locality, caught on leaf of Ulmus carpinifolia, 4.viii.1988, leg. R. Puplesis and V. Sruoga (MRL). Leaf-mines (no type material): Mangyschlak Region, Beyneu; Kara-Kalpaks- kaya ASSR, Kara-Kalpakiya. Diagnosis. - This species is similar to the Euro- pean B. ulmella Zeller (Seksjaeva 1981) (on Quer- cus) and the neartic B. electa Braun which also feeds on Ulmus leaves (Braun 1963). B. caspica is easily recognized by its male genitalia with a long vinculum, the presence of a transtilla and the form of the valvae. Description Male (fig. 4). - Forewing length 2.9-3.1 mm. Face creamy white, tuft brown or brownish in cen- tral part, with creamy white piliform scales later- ally. Eye-caps creamy white. Antennal flagellum with alternating white and brown rings of equal width. Thorax creamy white, slightly mottled, as some scales have brownish tips. Forewing with same colour as thorax, but some brown spots are present. Three or four of these spots are situated on the costal margin, the two below the fold, in apical part of forewing are smaller and usually darker. They include patches of blackish-brown scales. Dark-tipped scales form a line on the pale creamy cilia. Hindwings and cilia greyish cream to pale brownish. Legs cream with little fuscous shad- ing. Female. - Forewing length about 3.3 mm. Sim- ilar to male, but face cream, tuft slightly brownish centrally. The spots on the forewing may be larger than in male. Male genitalia (fig. 5). - Valva with convex mar- gin medially, suddenly tapering apically. Many large and some short setae on apical and medial part of valva. Transtilla present. Anellus forming an asymmetric ring. Uncus with two large lateral PUPLESIS ET AL.: Leaf-mining Lepidoptera setosae lobes. Vinculum ventrally a large, more or less triangular lobe, rounded anteriorly. Aedeagus long and slightly bent basally. Female genitalia (fig. 6). - Apophyses posteri- ores long. Ductus bursae narrow and very long. Bursa copulatrix more or less oval with numerous small spines. Egg. - Laid on the underside of a leaf of Ulmus carpinifolia Rupp. ex Suckow, generally close to the midrib or a lateral vein. In some cases, eggs are laid away from veins. Mine (fig. 8). - Gallery very narrow and compar- atively long (18-20 mm) with black linear frass, leaving clear margins throughout its whole course. Cocoon (fig. 7). - Whitish cream with blackish perpendicular patterns. Distribution (fig. 1). - Probably widespread in northern Caspiya, but possibly absent from the Amudar’ya oases and Turkmeniya. Gracillaridae Phyllonorycter schreberella (Fabricius) (fig. 9) Diagnosis. — It belongs to the Phyllonorycter ulmifoliella group, which larvae are usually leaf- miners of Ulmus (Kuznetzov 1981). It is easily recognized from all other species in that group by the silvery shining frons, thorax and base of the forewings. Tuft on head black. Forewing with two silver fasciae basally and medially, two silver spots apically. There is a black spot on base of forewing, near costal margin. Male genitalia (fig. 9) differs from all other species of the genus by the narrow and bent valvae. Distribution (fig. 1). - P. schreberella is found in Europe from England and Scandinavia to the Bal- kans and the European part of the USSR, also in Asia Minor, the Caucasus and the mountains of Turkmeniya (Kopet-Dag ridge) (Kuznetzov 1981). The species is here recorded for the first time from the northern Caspiyan region. Material examined. - 2 6, 1 9, USSR, Astrakhan re- gion, Baskuntschak, mines 4.viii.1988, ex p. 7.viii.1988, leg. R. Puplesis and V. Sruoga (MRL). ACKNOWLEDGEMENT We thank Zigmantas Gudzinskas (Vilnius) for the identification of Ulmus carpinifolia. 71 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Ke 3 : == ven ESS Ny . N ae EN yy Le E YW ù Y . RQ U U N VEN } si Fig. 4-7. Bucculatrix caspica. - 4, forewing (scale 1 mm); 5, male genitalia, holotype (scale 0.1 mm); 6, female genitalia, paratype (scale 0.1 mm); 7, cocoon, from type locality. 72 Fig. 8. Mine of Bucculatrix caspica on Ulmus carpinifo- lia. REFERENCES Braun, A., 1963. The genus Bucculatrix in America north of Mexico. - Memoirs of the American Entomological Society 18: 1-207. Johansson, R. & E. S. Nielsen, 1990. Tribus Nepticulini. - In: R. Johansson et al., The Nepticulidae and Opos- tegidae (Lepidoptera) of North West Europe. - Fauna Entomologica Scandinavica 23: 111-238, plates. Kuznetzov, V. I., 1981. Gracillariidae (Lithocolletidae) — Moli-pestrianski. - Opredelitel nasekomyh Evropeys- koy casti SSSR, Leningrad 4(2): 274, 277-278. PUPLESIS ET AL.: Leaf-mining Lepidoptera Fig. 9. Male genitalia of Phyllonorycter schreberella (F.) (scale 0.1 mm). Seksjaeva, S. V., 1981. Bucculatricidae - Krivousiye kro- hotki-moli. - Opredelitel nasekomyh Evropeyskoy casti SSSR, Leningrad 4(2): 136-148. Tushinskij, G. K. & Davydova, M. I., 1976. Fiziceskaya geografiya SSSR. - Prosvesceniye, Moscow: 179. Received: 5 january 1990 Revised version accepted: 22 November 1990 73 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 BOOK REVIEWS Emmet, A. M., 1991. The scientific names of the British Lepidoptera. Their history and meaning. - Harley Books, Colchester, England. 288 pp., viii plates. [ISBN 0 946589 35 6, paperback edition. Price ca US $ 40]. Many entomologists consider scientific names simply as labels of the subjects of their studies, and do not bother about their meaning. Others, how- ever, are interested in the history of the name and wonder why they are named as they are. To cite the author of this book: ‘There is no need to seek the explanations of scientific names, just as there is no need to do crossword puzzles. There are, however, many who derive pleasure from both pursuits.’ For those who are interested in the history of names in general and nomenclature of Lepidoptera in particular, Colonel Emmet has provided a wealth of interesting information, with explana- tions of names of all British species, genera and families, including a few synonyms, 3000 names in total. The author not only tried to give the exact meaning of the latin or greek words from which the name supposedly was derived, but also tried to trace the reasons of the author for giving it this particular name. This was no small task, when one realises that very few authors before this century ever explained the new names they were propos- ing. In many cases therefore Emmet had to guess at the thoughts of the name-giving authors, but was greatly helped by his immense knowledge of British Lepidoptera. The systematic section of this book therefore, is by no means a dry dictionary, but well readable and full of small interesting stories about authors of the past. Emmet also could rectify many earlier explanations, by his combined know- ledge of the classic languages and Lepidoptera. That with so many names a few misinterpreta- tions or errors were noted, is a very minor flaw indeed. To cite some, noted by the reviewer: 34. Ectoedemia occultella. It is very unlikely that Lin- naeus was indicating the larval habits of this spe- cies: he described it from specimens which he found on his window. The genus Trifurcula (page 45) was named after the three-forked radius in the hindwing, not forewing. Caradja (species 871a) was a Rumanian entomologist, not German. The introduction is a well written history of nomenclature in Lepidoptera, and gives insight in the way early authors, like Linnaeus, formed scien- tific names. It also explains the curious usage of special endings for particular groups of Lepidop- tera (such as -ella, -ana, -ata). Emmet follows, 74 against his classical background, the current lepi- dopterological practice to use original spellings and endings, without bothering about gender of the generic name. Colonel Emmet has written an invaluable source of information and fun. The book is warmly recom- mended to all entomologists interested in names, and should not be missing in the library of any lepidopterist. [E. J. van Nieukerken] Rubtsov, I. A., 1990. Blackflies (Simuliidae). - Fauna of the USSR, Diptera, Volume 6, Part 6: p. i-xxviii + 1- 1042, figs. 1-424 [mostly composite]. [ISBN 90 04 088717, distributed outside India by E. J. Brill, Leiden. Price 300 Dutch Guilders, appr. us $ 150]. According to information provided by the pub- lishers ‘the purpose of the present volume is to provide brief descriptions of species and new iden- tification keys. It primarily incorporates numerous additions to the first addition of the Fauna of the USSR. This second edition also includes 18 species from countries adjoining the Palearctic region, which have not been recorded to-date in the Soviet Union, and 30 species described by Enderlein from Europe (whose description has been improved upon), which may be discovered later in the Soviet Union’. Usually translations of Russian handbooks are most welcome, but one may doubt whether this is also true in this particular case. Firstly the original was published as early as 1956. Secondly, the text is clearly out of date, since a revised and more comprehensive work by the same author appeared between 1959 and 1964 in Lindner's Die Fliegen der palaearktischen Region’ in the German lan- guage. Thus, for scientists who understand Ger- man the information was already available for more than 25 years. I suppose that most dipterists not fluent in German or Russian would have pre- ferred a translation of Lindner rather than one of the Fauna of the ussr. The translation as such seems to be done with reasonable care. For in- stance, the translator notifies the reader about pos- sible mistakes in the original text, and there is also a translation of the lettering in the illustrations. As usual the quality of the Indian printing is far behind European standards, making some figures difficult to understand. Comparing the three works it ap- pears that the figures in Lindner are by far the best. [J. van Tol] BERNHARD]. VAN VONDEL Natuurmuseum Rotterdam REVISION OF THE PALAEARCTIC SPECIES OF HALIPLUS SUBGENUS LIAPHLUS GUIGNOT (COLEOPTERA: HALIPLIDAE) Vondel, B. J. van, 1991. Revision of the palaearctic species of Haliplus subgenus Liaphlus Guignot (Coleoptera: Haliplidae). - Tijdschrift voor Entomologie 134: 75-144, figs. 1-312. (ISSN 0040-7496]. Published 1 July 1991. The palaearctic species of the subgenus Liaphlus Guignot, genus Haliplus Latreille, are revised. Thirty species are recognised. The following new synonyms are established : H. modestus Zimmermann as a junior synonym of H. eximis Clark, H. subguttatus Crotch and H. salinarius Wallis of H. fulvus (Fabricius), H. pelopis Sahlberg of H. guttatus Aubé, H. syriacus Wehncke and H. zimmermanni Gschwendtner of H. maculatus Motschulsky and H. transvolgensis Semenov of H. variegatus Sturm. Eight new species are described: H. angusi, H. astrakhanus, H. davidi, H. excoffieri, H. gafnyi, H. holmeni, H. jaechi and H. ortali. A key to the species is provided. Most primary types have been examined. Lectotypes have been designated for H. abbreviatus Wehncke, H. andalusicus Wehncke, H. basinotatus Zimmermann, H. dalmatinus Miiller, H. dalmatinus var. weberi Miiller, H. interpunctatus (Marsham), H. lapponum Thomson, H. fulvus sparreschneideri Muns- ter, H. guttatus Aubé, H. maculatus Motschulsky, H. pelopis Sahlberg, H. cinereus Aubé, H. ovalis Sharp, H. sharpi Wehncke, H. syriacus Wehncke, H. zimmermanni Gschwendtner, H. variegatus ab. pallidior Müller, H. leopardinus Sahlberg and H. trans- volgensis Semenov. B. J. van Vondel, Roestuin 78, 3343 CV Hendrik Ido Ambacht, The Netherlands. Key words. - Liaphlus, palaearctic region, China, new species. CONTENTS species have been described from the southern and eastern parts of the region. Various problems in INCIOGUGHONEL E TRE I DCR dé dent ia e OL cence: MEEO a nn 76 sa 8 RESSE SYSTEMALICISECLIO DEEE EE CE A 77 Le noe ESTER IEN ChecklisttofitreatedispeCie Sto Rn 78 Da Ee Key to the palaearctic species of Liaphlus … 78 Description of the palaearctic species O2 DUI assi eri e 81 Unnamed material ............................... 135 Acknowled SEMENTI 125 RÉ LÉLENICES nr AR it Re LE hr 136 MAPS TE I I 140 INTRODUCTION This revision deals with the palaearctic species of Liaphlus, a subgenus of Haliplus, one of the five genera of the waterbeetle-family Haliplidae. The latest revision of Haliplus in the palaearctic region was by Zimmermann (1924). The status of some species remained uncertain. Afterwards some new for the Haliplus-species lacking the pronotal basal plicae and possessing a setiferous striole on the dorsal side of the metatibia. Afterwards Guignot (1955) proposed a division of Liaphlus into 9 spe- cies-groups, of which 4 groups are represented in the palaearctic region as treated in this revision. Especially outside the palaearctic region his divi- sion is certainly in some cases based upon misin- terpretations. For instance the oriental species Ha- liplus angustifrons Régimbart and H. arrowı Guignot are placed in different groups. In my opin- ion however, these species are very closely related. A similar case is Haliplus pulchellus Clark and H. diopus Guignot, also placed in different groups, but also closely related. The subgeneric status of Haliplus laminatus (Schaller) is not completely clear. According to the 75 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Fig. 1. Haliplus flavicollis, dorsal view. - an, antenna; as, apical spur; cl, clypeus; el, elytron; fr, frons; 1, 2, 3, etc, first, second, third, etc. elytral interval; p, primary puncture-row; pr, pronotum; s, secondary puncture-row; su, suture; ss, setiferous striole; t, tibia; ve, vertex. Fig. 2. Haliplus flavicollis, ventral view. - cp, hind coxal plate; ee, elytral epipleuron; mp, metasternal process; pe, proepisternum; pp, prosternal process; ps, prosternum. definitions for Liaphlus it should belong to that subgenus, but the larval stages are distinctly differ- ent and very much like the larvae found in the subgenus Haliplus s. str. (sensu Guignot 1928) (Vondel 1986). Beutel & Ruhnau (1990) made a phylogenetic analysis of the genera of Haliplidae based on char- acters of adults. However, on subgeneric level and below much more study is needed to elucidate the phylogenetic relationships within the Haliplidae. For that reason I refrain from dividing the species in this revision into species-groups. They are simply treated in alphabetical order. For many species the type-material was studied. For some species, of which the status seemed clear to me, only few or no attempts have been made to trace and study the type-material. In some other cases several attempts to locate the types failed. About 2800 specimens have been studied, mostly from Western Europe. Eastern and Southern pa- laearctic material is relatively scarce in the collec- tions I had access to. 76 For all species (re)descriptions are provided. Female genitalia are not described, because they usually are very uniform and further research is necessary to check if there are characters reliable enough to separate related species. Descriptions of larvae are not included. METHODS Before dissecting the aedeagus the beetles are softened in water with detergent for about 2 hours. The aedeagus is pulled out with a small hooked needle and placed into a drop of alcohol. During dissection the beetle has to be kept wet, to prevent the severed aedeagus jumping out of sight. In alco- hol the left- and right parameres are separated from the penis by using fine needles. When the alcohol has almost dried up the three parts are usually glued with Fauré-mixture on the same piece of cardboard as the beetle. The penis and the par- ameres are positioned so that they are seen from the side. The water-soluble Fauré-mixture makes it possible to loosen the genitalia to study them in an other position or to make temporary slide mounts. In some cases the genitalia are transferred into Euparalona plastic card, covered by a coverslip and pinned to the original needle. Genitalia should not be glued to plastic cards with water-soluble glue as they are liable to become detached in time. Drawings of genitalia are made by using a Swift universal microscope with a camera lucida attach- ment. The other drawings are made by using a Zeiss binocular with an ocular-micrometer. Measurements are made by using the ocular- micrometer of a Zeiss binocular. Morphological terms are according to figs. 1 and DA _ The palaearctic region as treated in this revision comprises Europe including Iceland, North Africa north of the Sahara and Asia north of the line formed by the northern borders of Pakistan, India, Nepal, Bhutan, Burma, Laos and Vietnam. For practical reasons Taiwan and China as a whole are taken into account, although the south-eastern part has an oriental fauna. Material of many institutions and private per- sons has been studied, including a number of prim- ary types. Locality-names are given, as far as pos- sible, according to the Times Atlas of the World (Comprehensive edition, 1983). Deviating names on original labels are cited, when necessary, in parentheses. In a number of cases I was not able to trace the locality, because of the poor or unreadable labels. Countries are used with their present-day boundaries. Distribution-maps are based on material exam- ined. Reliable literature-records are included, but using different symbols. Data on biology are seldom present on labels, so notes on the biology are mainly based on literature and the author's experience. If nothing is menti- oned in the descriptions about biology, such infor- mation was unknown to the author. The institutions from which material has been examined or which are otherwise referred to in the text have been abbreviated according to Arnett & Samuelson (1986) with the addition of: HUJI - IES Laboratory, Hebrew University of Jer- usalem, Israel. MSNT - Museo Civico di Storia Naturale, Trieste, Italy. MOG - Museum voor het Onderwijs Museon, 's Gravenhage, Netherlands. RIN — Rijks Instituut voor Natuurbeheer, Leersum, Netherlands. Further material from the following private col- lections has been studied: Angus, Englefield Green, VAN VONDEL: Palaearctic Liaphlus UK; van Berge Henegouwen, Zoetermeer, Ne- therlands; Cuppen, Ede, Netherlands; Drost, Wadenoijen, Netherlands; Giessen, Gaanderen, Netherlands; Hielkema, Gouda, Netherlands; Huijbregts, Leidschendam, Netherlands; Jongema, Wageningen, Netherlands; Kanaar, Leiderdorp, Netherlands; Leblanc, Troyes, France; Matsui, Hondo City, Japan; Onder de Linden, Krimpen aan de IJssel, Netherlands; Rotzer, Gampel, Switzer- land; Schilthuizen, Leiden, Netherlands; Schreijer, Alkmaar, Netherlands; Sung Hwa Lee, Taegu, Ko- rea; Vallenduuk, Lelystad, Netherlands; Vondel, Hendrik Ido Ambacht, Netherlands. SYSTEMATIC SECTION Subgenus Liaphlus Guignot Haliplus sg. Liaphlus Guignot, 1928: 138. Type species: Dytiscus fulvus Fabricius, 1801, by subsequent desig- nation (Guignot 1930: 75). Haliplus sg. Hoplites Kinel, 1929: 219. Preoccupied. [See remarks under Haliplus laminatus.| Type species: Dy- tiscus laminatus Schaller, 1783, by monotypy. Liaphlus Guignot; Guignot 1933: 228, 1947: 48, 1959: 33, Balfour-Browne 1936: 76, 1938: 12, Csiki 1946: 558, Zaitsev 1953: 43, Freude 1971: 15, Galewski 1976: 27, Franciscolo 1979: 108, Vondel 1986: 132, Holmen 1987: 110, Beutel & Ruhnau 1990: 11. Diagnosis. - The following character set is diag- nostic for specimens of this subgenus: 1. Pronotum without basal plicae. 2. Metatibia with a setiferous striole on dorsal face. 3. Males: right paramere with a solid digitus. This is not the case in H. laminatus (see remarks under that species). Remarks. - Liaphlus is represented in all faunal regions. Beutel & Ruhnau (1990) examined a large number of characters of nine species of Liaphlus from different faunal regions, but they did not find evidence for the monophyly of the subgenus. Within the Haliplidae Liaphlus can be recog- nised by means of the following keys. Key to the genera of world Haliplidae 1. Last segment of palpi longer than penultimate segment. Metacoxal plates at least covering part of sixth abdominal sternite................. enna SV Peltodytes Régimbart — Last segment of palpi clearly shorter than pen- ultimate segment. Last three abdominal ster- mites sreely visible: a. RR 2 VY TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 2. Head as wide as pronotum. Elytra strongly PUNCLULEKATORMALOWSEL a. We ARES AS PRESS … Algophilus Zimmermann - Head not as wide as pronotum. Elytra with GIEATRFOWSTO{IPUNELULESPRBRARA RE E 3 3. Pronotum almost square, front-corners strongly rounded, elytra usually with longitud- INA ALES ARE RENEE Brychius Thomson — Pronotum widest at base, sides converging an- teriorly, elytra without longitudinal ridges in ÉirsusinINtervals PE MNT ES ETAPE STO 4 4. Median part of prosternum and base of pro- sternal process forming a plateau-like eleva- tion, at least in part angularly separated from sides of prosternum......... Haliplus Latreille - Prosternum evenly rounded from side to side SONA RE Sc Apteraliplus Chandler Note: Algophilus is restricted to South Africa, while Apteraliplus is only known from North America. Key to the palaearctic subgenera of Haliplus: 1. Dorsal and ventral side of body with clearly visible (30 x) micropuncturation, on elytra be- tween small primary punctures.................. LR INI IA DIREI Haliplidius Guignot — Elytra smooth or with much weaker micro- puncturation (visible at 50 x) between strong PEIMaryapUneture OWS et ere 2 2. Hind tibia with a setiferous striole on dorsal face. Pronotum without basal longitudinal pli- Carel, Lie Meng Liaphlus Guignot - Hind tibia without a setiferous striole on dorsal face. Base of pronotum on both sides with a, sometimes very short, rarely absent, longitud- inaljplicamiparzi na: st ren em. 3 3. Pronotal plicae reaching beyond middle and strongly bent.......... Neohaliplus Netolitzky — Pronotal plicae not reaching beyond middle, straight or slightly curved..... Haliplus s.str. Checklist of palaearctic Haliplus (Liaphlus) 1. H. abbreviatus Wehncke, 1880 2. H. andalusicus Wehncke, 1874 3. H. angusti sp. n. 4. H. astrakhanus sp. n. 5. H. basinotatus Zimmermann, 1924 6. H. chinensis Falkenström, 1932 7. H. dalmatinus J. Miiller, 1900 8. H. davidi sp. n. 9. H. diruptus Balfour-Browne, 1946 0. H. excoffieri sp. n. — 78 30. . H. eximis Clark, 1863 H. modestus Zimmermann, 1924 syn. n. H. flavicollis Sturm, 1834 H. fulvus (Fabricius), 1801 H. subguttatus Crotch, 1873 syn. n. H. salinarius Wallis, 1933 syn. n. . H. gafnyi sp. n. . H. guttatus Aubé, 1836 H. pelopis Sahlberg, 1907 syn. n. H. holmeni sp. n. . H. jaechi sp. n. H. kotoshonis Kano & Kamiya, 1931 H. kulleri Vondel, 1988 H. laminatus (Schaller, 1783) . H. maculatus Motschulsky, 1860 H. syriacus Wehncke, 1880 syn. n. H. zimmermanni Gschwendtner, 1921 syn. n. H. maculipennis Schaum, 1864 H. mucronatus Stephens, 1828 H. ortali sp. n. H. ovalis Sharp, 1884 H. pulchellus Clark, 1863 H. rubidus Perris, 1857 H. sharpı Wehncke, 1880 H. variegatus Sturm, 1834 H. transvolgensis Semenov, 1904 syn. n. H. villiersi Legros, 1972 Key to the palaearctic species of Liaphlus . Distance between eyes about eye-width (fig. 1705208244) een ee RE 2 Distance between eyes at least 1.2 X eye- wıdchs3e. au. ali anna ere ee 4 . Prosternal process furrowed on both sides (fig. DASEIN TR 26.pulchellus Prosternal process strongly impressed in mid- dle, at least in apical half (fig. 174)............ 3 . Elytra with basal black band, eighth puncture- row with about nine small dark blotches....... ee brite. corte sestri. A 22.maculipennis Elytra without basal black band, eighth punc- ture-row not with dense row of dark blotches zeide arie see PRIE 18.kotoshonis . Head relatively wide (fig. 217), distance be- tween eyes at least 2.3 X eye-width. Metaster- nal process flat in middle. Elytra without dark blorches sh pere 23.mucronatus Head narrower. Metasternal process at least with small pit or groove in middle........... 5 . Sides of prosternal process before coxae ex- tending forwards as complete plicae to anterior edge of prosternum (LR) re eer 8 Sides of prosternal process before coxae not completely extending forwards as plicae to an- 10. Male 12. terior edge of prosternum (fig. 116).......... 6 . Plicae on sides of prosternal process widely interrupted before coxae (fig. 17). Longer ap- ical spur of hind tibia usually as long as or longer than first tarsal segment (fig. 15). Elytra mostly with darker blotches..... 2.andalusicus Plicae on sides of prosternal process com- pletely absent before coxae (fig. 116). Elytra without clear blotches. Apical spurs of hind tibia not as long as first tarsal segment....... N . Punctures of primary elytral rows wide in basal area, distance between rows not more than diameter of punctures (fig. 255). Distance be- tween eyes 1.7-1.9 X eye-width..... 27.rubidus Punctures of primary elytral rows rather small, distance between rows more than diameter of punctures (fig. 113). Distance between eyes ASG) eye width deli oa 12.flavicollis . Setiferous striole on dorsal face of hind tibia stretched over apical half (fig. 195). Elytra us- ually with more or less confluent and inter- rupted dark lines following rows of dark punc- tures, but distinct blotches usually absent. Basal elytral punctures coalescing to form short grooves (fig. 188). Male: first tarsal segment of midlegs strongly dilated ventrally (fig. 192). Female: elytra almost completely covered with MICrO-PUNELUTESE te ee 20.laminatus Setiferous striole on dorsal face of hind tibia usually not longer than % X tibia length, not reaching apical point. Male: first 3 tarsal seg- ments of midlegs weakly dilated ventrally. Female: elytra without clear micropunctua- OR IA Ae bore brede 9 . Elytra without dark blotches. Pronotal and ely- tralipunctures black terme tee ll.eximis Elytra with dark blotches, or when these blotches are strongly reduced or absent punc- tures on disc of pronotum are not black ....10 Elytra with basal transverse black band as dark asisuturalimaculation #7 e. 34 Elytra without basal black band. Sometimes some transverse darkening separated from ba- sal margin or a brown basal band, clearly less dark than sutural maculation.................. 11 Widened basal punctures on pronotum twice the width of elytral primary punctures (fig. 141). Body-shape parallel or subparallel. Pro- sternal process narrow, length 2.6 X width, slightly narrowed near coxae (fig. 145). Elytra with mostly vague blotches, one large blotch on middle of suture and three smaller ones on CACHE ME ONE RARO 15.guttatus Basal punctures on pronotum usually not twice as wide as elytral primary punctures. Body not clearlygparallel ee ea 12 Weng ches CLS Inn RIINA PITTORI 29 16. We . Metasternal VAN VONDEL: Palaearctic Liaphlus Eng: 13 . Elytra with longitudinal distinct blotches in even intervals, blotches not connected to dar- kenedisutune stp s-te oe eden nale 14 Elytraliblotchesidifferent +... sr... 17 . Metasternal process flat anteriorly. Vertex dar- kenedi4& eee azar bea 15 Metasternal process grooved anteriorly. Ver- tex not darkened . Setiferous striole weak, about Y, X tibia length (fig. 33). First elytral primary puncture-row with about 28 punctures. Pronotum anteriorly partly bordered. Males: left paramere with solid digitus (fig. 36). at es. 4.astrakhanus Setiferous striole strong, about % X tibia length (fig. 202). First elytral primary punc- ture-row usually with more than 28 punctures. Pronotum anteriorly not bordered. Males: left paramere without solid digitus.................. specimens of 21.maculatus Distance between eyes less than 1.7 X eye- width. Males: top of penis short (fig. ON er specimens of 7.dalmatinus Distance between eyes more than 1.7 X eye- width. Males: top of penis long, usually slightly widenedkapicallya(fig 278) ee 2... ee REN es specimens of 29.variegatus Elytra with large maculation, discal blotch and blotch in apical part connected to darkened suture sl ae chaos. tases eke ER 18 Elytra with usually vague maculation on first and even intervals, or only darkening on discal parts of puncture-rows, blotches on apical part Nnotconnected to suture: ar. sees oe 21 . Length of prosternal process 2.6 X width (fig. ZI desde EER 3.angust Length of prosternal process less than 2.4 X width: mite Aree Er ea. 19 . Prosternal process hardly narrowed near coxae (fig. 73). Males: left paramere with short dig- IUSR 8.davidi Prosternal process clearly narrowed near coxae. Males: left paramere without digitus 20 . Distance between eyes more than 1.7 X eye- Width en specimens of 29.variegatus Distance between eyes less than 1.7 X eye- wide oon specimens of 9.diruptus process grooved anteriorly, weakivyapunctured nace: tener tes le. Sens 22 Metasternal process flat anteriorly........... 23 . Prosternal process impressed posteriorly (fig. 287). Elytral primary puncture-rows sparse, sutural punctures weak (fig. 285) 30.villiersi Prosternal process posteriorly at most with weak transverse impression (fig. 7). Elytral primary puncture-rows dense, sutural punc- turesistrongd(fig-)) hanna. l.abbreviatus 79 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 © © GOOG © | | | “| | © Cee Sed è d è 80° 5 oe AL © 0 0 0 e o 0 e o Oe , locality unknown. - 3, dorsal view; 4, antenna; 5, elytral punctuation; 6, dorsal side ; 7, prosternal process; 8, lateral view of prosternal process; 9, left paramere; 10, penis; 11, right paramere. 80 Figs. 3-11. Haliplus abbreviatus of hind tibia; DIE 28. 29} 30. 31. 32. WE Malest SEWER EI RER Silent EHE tie suli 24 Female sso ed e ona 27 24. Left paramere without solid digitus 14.gafnyz Left paramere with solid digitus.............. 25 . Digitus on left paramere long and parallel- sided. Base of pronotum with about five stronger punctures on both sides.... 19.kulleri Digitus on left paramere tapering. Base of pronotum with almost continuous row of SCLONDEM PUNGCULES AR ee. 26 . Left paramere with strongly developed taper- ing digitus, row of long hairs on inner margin Ofiparamere Kir ne Oars. ere L 17 jaecht Left paramere with small tapering digitus, at most some short hairs on inner margin of Paramere te ois. MESA PE LA ANN! 24.ortali Metasternal process flat with shallow pit in middle, strongly and densely punctured, small punctures between large ones (fig. 136)........ 14.gafnyi Metasternal process with clear pit in middle, sparsely and moderately strongly punctured EEE WEES ER RENT IO NAME, OU 28 Basal punctures on pronotum weak and re- duced in the middle (fig. 179). Prosternal pro- cess weakly narrowed (fig. 183)..... 19.kulleri Base of pronotum with continuous row of strong punctures (fig. 161). Prosternal process Stronplysnarrowed! (fig IGS e 17.jaechi or 24.ortali Distance between eyes less than 1.4 X eye- WALTER A RAET, OPCs LIL OI 2058 30 Distance between eyes more than 1.4 X eye- Wit ch MEREN epr: Je SMOLIN RICE, 31 Black suture not reaching first secondary punc- ture-row in basal part (fig. 91). Body strongly tapering apically. Penis and parameres figs. 99- NOME SOR EEE A A MEE BAM 10.excoffieri Black suture reaching first secondary punc- ture-row in basal part (fig. 48). Body more oval. Penis and parameres figs. 54-56.......... SRO SO NI 6. chinensis Elytral discal blotch connected to suture. Penis and parameres figs. 241-243........... 25.ovalis Elytral discal blotch not connected to su- EUDORA ee een ne 32 Length of hind tarsus including claws more than 1.3 X tibia length. Prosternal process slightly narrowed near coxae, narrowed part 0.6 to 0.7 X widest part. Punctures of setiferous striole not connected in a groove. Male: digitus of right paramere 1 X length of basal part.... large specimens of 7.dalmatinus Length of hind tarsus less than 1.3 X tibia length. Prosternal process strongly narrowed near coxae, narrowed part 0.5 X width of wid- ESCHPALELIIR sae een 33 33: 34. 39) 36. VAN VONDEL: Palaearctic Liaphlus Distance between eyes less than 1.6 X eye- width. Setiferous striole on dorsal side of hind tibia not strongly grooved, separate points us- ually clearly visible. Male: apical part of penis very narrow and pointed (fig. 205)............. 21.maculatus Distance between eyes more than 1.6 X eye- width. Setiferous striole on dorsal side of hind tibia strongly grooved, separate points not or hardly visible. Male: apical part of penis wide and rounded at the top (fig. 127).....13.fulvus Length 3.5-4.1 mm. Black sutural stripe in an- terior half at most reaching first secondary PUNCEUIE LOWER RS 5.basinotatus Length at most 3.6 mm. Black sutural stripe reaching first primary puncture-row......... 35 Distance between eyes about 1.2 X eye-width. Prosternal process wide, impressed apically, weakly narrowed near coxae (fig. 155). Setifer- ous striole on dorsal side of hind tibia consist- ing of about 6 isolated points, not grooved (fig. 154). Male: left paramere with solid digitus (fig. 158). Between fore-claws a small cushion (OTR DS de A o al 16.holmeni Distance between eyes 1.3-1.5 X eye-width 36 Fourth and fifth antennal segment about as long as wide (fig. 263). Punctures in first to fourth primary elytral rows weaker than in other puncture-rows. Prosternal process broad, hardly narrowed near coxae, hardly or not impressed apically (fig. 266). Anterior edge of prosternum clearly and completely bor- RATER eme er TIERRA 28.sharpt. Fifth antennal segment clearly longer than wide. Inner primary puncture-rows of elytra not weaker than outer ones. Prosternal process strongly narrowed near coxae, strongly im- pressed apically. Anterior edge of prosternum only bordered near process. South-east pa- laearctic or north oriental specimens OPOLE, NERE TT 9.diruptus DESCRIPTION OF THE PALAEARCTIC SPECIES OF LIAPHLUS 1. Haliplus abbreviatus Wehncke (figs. 3-11) Haliplus abbreviatus Wehncke, 1880: 73. Lectotype 9 (here designated), ‘Mesopotamia, Sharp’ (MNHN) [examined]. [Iraq] Haliplus abbreviatus; Zimmermann 1920: 303, 1924: na 136, Alfieri 1976: 38. Diagnosis This species is closely related to villzerst or even conspecific. Specimens of vzlliersi have the proster- | process more impressed posteriorly, the prim- 81 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 | 08666 ® Figs. 12-20. Haliplus andalusicus, lectotype. - 12, dorsal view; 13, antenna; 14, elytral punctuation; 15, dorsal side of hind tibia; 16, prosternal process; 17, lateral view of prosternal process; 18, left paramere; 19, penis; 20, right paramere. 82 ary elytral puncture-rows sparser and the sutural punctures weaker. Description Length 3.0 mm, width 1.7 mm. Body oval, widest in the middle (fig. 3). Head. - Light brown, rather strongly punctured, more sparsely punctured between the eyes (fig. 3). Distance between the eyes 1.6 X eye-width. Anten- nae (fig. 4) and palpi yellow-brown. Pronotum. - Yellow-brown, base and front mar- gin slightly darkened. Lateral borders straight to slightly convex, with fine dark margins. Disc spar- sely punctured, front margin behind an unpunc- tured zone with a strongly punctured band, base strongly punctured, basal puncture-row partly wi- dened and darkened (fig. 3). Elytra. - Yellow-brown with in the anterior half fairly strong and apically weaker primary punc- ture-rows, punctures brown to almost black (fig. 5). First primary puncture-row with about 30 punc- tures. Secondary puncture-rows fine. Brown, some- times vague markings on the even intervals, on the apical point and along the apical % of the suture, median spot connected to the suture. Ventral side. - Body yellow-brown to yellow-red, prosternal process partly dark-brown, elytral epi- pleura yellow, legs yellow-brown with some dar- kening around the coxae and trochanteres. Proster- nal process flat, slightly impressed apically, narrowed near the front coxae, strongly punctured (fig.7), laterally completely bordered (fig.8). Meta- sternal process furrowed, deep pit in the middle, strongly punctured (fig.7). Setiferous striole on dorsal face of hind tibia with about 6 punctures, longer of the two tibial spurs almost as long as the first tarsal segment (fig.6). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 9-11. Distribution (fig. 305) Iraq, Iran?. Alfieri (1976) gives Egypt. Remarks This species is described here on the basis of a single female (lectotype) and two male specimens, which are considered to belong to this species (figs. 3-11). It is probably a very variable species, which often may be confused with variegatus. There is a good chance that a number of the middle eastern specimens, in literature cited as variegatus belong to abbreviatus. Material examined. - 3 ex. - Iraq: 1 Q (lectotype). - Iran(?): 1 6, Turcom, Kurd., 15943, Millingen, Fry Coll. (BMNH) — Without locality: 1 @, Reitter (NHMW). VAN VONDEL: Palaearctic Liaphlus 2. Haliplus andalusicus Wehncke (figs. 12-20) Haliplus andalusicus Wehncke, 1874: 135. Lectotype & (here designated), [Spain] ‘Andalusia, Dieck, Dr. Guignot visité 1925’ (MNHN) [examined]. Haliplus andalusicus, Marseul 1882: 107, Seidlitz 1887: 30, Escalera 1914: 57, Scholz 1916: 17, Zimmermann 1920: 303, 1924: 136, Guignot 1933: 232, 1947: 52, 1959: 35, Burmeister 1939: 214, Lagar Mascaro 1968: 73, Hoch 1972: 251, Ienistea 1978: 294. Diagnosis This species is most easily confused with varte- gatus from which it can be distinguished by the interrupted lateral plicae of the prosternal process and the apical spur of the hind tibia being as long as or longer than the first tarsal segment. Description Length 2.5-3.0 mm, width 1.4-1.7 mm. Body oval, widest in the middle (fig. 12). Head. - Yellow-brown with usually brown ver- tex, sparsely punctured. Distance between the eyes 1.9-2.0 X eye-width. Antennae (fig. 13) and palpi yellow to yellow-brown. Pronotum. - Yellow to yellow-red. Lateral borders straight to slightly convex, finely bordered. Sparsely to rather strongly punctured, basal punc- tures slightly widened and darkened (fig. 12). Elytra. - Yellow to yellow-red, primary punc- ture-rows rather weak with exception of the stronger punctures in the anterior half of the first three rows, 32-36 punctures in the first row (fig. 14). Secondary punctures weak. Darkening of the punctures not always visible. Brown to almost in- visible markings on the even intervals, on the ap- ical point and along the apical % of the suture, the median spot may be connected to the suture. Ventral side. - Body yellow to yellow-brown, elytral epipleura yellow, legs yellow-brown. Pro- sternal process impressed in the middle, strongly punctured, narrowed near the coxae (fig. 16), lat- eral plicae interrupted before the coxae (fig. 17). Metasternal process grooved with a pit in the mid- dle to largely impressed, sparsely punctured (fig. 16). Setiferous striole on dorsal face of hind tibia with 4-8 punctures, longer of the two tibial spurs as long as or longer than the first tarsal segment (fig. 15). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 18-20. Biology In fresh and especially brackish stagnant water in pools near the coast and more rarely inland. 83 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 21-29. Haliplus angusi, holotype. - 21, dorsal view; 22, antenna; 23, elytral punctuation; 24, dorsal side of hind tibia; 25, prosternal process; 26, lateral view of prosternal process; 27, left paramere; 28, penis; 29, right paramere. 84 Distribution (fig. 298) South of France, Spain (Burmeister 1939, gives The Baleares), Morocco, Algeria, Tunisia. Material examined. — 38 ex. - France: 1 9, La Capelle [dep. Lozère], 30.vi/iv.1987 (Leblanc); 6 ex., Albaron, Camargue, A. Chobaut; 1 ex., Montpellier, Et. Laragne, V.M. Duchon (MNHN). — Spain: 1 @ (lectotype); 1 & (paralectotype), Malaga, Sammlung Cl. Müller (zsMC); 1 ex., Gibraltar; 1 ex., Palencia (BMNH). - Morocco: 4 8, Tanger, Rolph (BMNH). - Algeria: 2 ex., Massif des Mou- zaia (BMNH); 4 ex., Philippeville, A. Thery (NHMw, MNHN); 1 ex., Biskra, surroundings, de Vauloger; 1 ex., Biskra, Mertu[?]; 2 ex., no further data (NHMW).-Tunisia: 2 8, Gafsa (NHMW); 2 6, 1 9, 3 ex, surroundings Gafsa, de Vauloger (NHMW, MNHN); 2 ex., Southern Tunisia, ili.iv.1925, ex coll. J. Omer Cooper (BMNH).-Locality un- known: 2 @ (paralectotypes), ex. coll. Wehncke (MNHN). 3. Haliplus angusi sp. n. (figs. 21-29) Type material. - Holotype 6, USSR, Siberia, 28.v- 28.vi.1982, Novosibirsk oblast, Karasuk, 53.5 N, 78 E, R. B. Angus (ZMAS). Diagnosis This species can be distinguished from related ones by the very narrow prosternal process. Possi- bly there is material in collections representing this species, but identified as H. maculatus Mot- schulsky. Description Length 3.3 mm, width 1.7 mm. Body oblong, subparallel, widest in the middle (fig. 21). Head. - Yellow-red, dark mark on vertex, weakly and sparsely punctured. Distance between the eyes 1.6 X eye-width. Antennae yellow (fig. 22), palpi yellow. Pronotum. - Yellow to yellow-brown on the disc. Lateral borders straight, lateral margin clearly finer anteriorly. Strongly darkened, occasionally widened punctures along the base, anteriorly rather weakly punctured, otherwise almost un- punctured. Along the central part of the base slightly impressed (fig. 21). Elytra. - Pale yellow, primary puncture-rows moderately strong, about 33 punctures in the first row (fig. 23). Secondary punctures in the first in- terval almost as strong as the primary punctures, other secondary punctures rather weak, accompan- ied by small punctures (fig. 23). Extensive mark- ings: a large arrow-shaped blotch on the disc, on the suture and the apex, often connected oblong blotches in the even intervals. The basal margin, normally covered by the pronotum, yellow-brown till the fifth striae (fig. 21). VAN VONDEL: Palaearctic Liaphlus Ventral side. - Yellow to yellow-red, elytral epi- pleura pale-yellow, abdominal segment brown bor- dered, legs yellow with some darkening near the coxae. Prosternal process narrow, flat, narrowed near the coxae, coarsely punctured (fig.25), lateral plicae complete (fig. 26). Metasternal process flat with a pit in the middle, sparsely punctured (fig. 25). Coxal plates sparsely and weakly punctured, near the suture almost unpunctured. Setiferous striole on dorsal side of hind tibia over % of the tibia length, with about 10 punctures, longer of the two tibial spurs % X the length of the first tarsal segment (fig. 24). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 27-29. Etymology. - This species is named after the collector Dr. Robert Angus. Distribution (fig. 304) Only known from the type-locality: USSR, Sibe- ria, Novosibirsk oblast, Karasuk. 4. Haliplus astrakhanus sp. n. (figs. 30-38) Type material: Holotype 8, [ USSR, Astrakhan] ‘As- tracan, Koltze’ (MNHN, Wehncke collection). Diagnosis This species is closely related to ortalz, but in the males the top of the penis is narrower and more pointed. Description Length 3.4 mm, width 1.9 mm. Body oval, widest in the middle (fig. 30). Head. - Red-brown, weakly punctured, vertex little stronger punctured. Distance between the eyes 1.5 X eye-width. Antennae (fig. 31) and palpi yellow-red. Pronotum. - Yellow-red, slightly darker on the disc and along the base. Lateral borders straight, finely bordered. Anteriorly partly bordered. Width 1.8 X the length in the middle. The middle of the base slightly impressed. Weakly punctured, along the base a row of stronger not or hardly darkened punctures (fig. 30). Elytra. - Yellow-red, many longitudinal vague blotches on the even intervals and along the suture (fig.30). Primary puncture-rows moderately strong, about 28 punctures in the first row. Secon- dary punctures weak, accompanied by very small ones (fig. 32). All punctures darkened. Ventral side. - Yellow-red to red-brown, elytral epipleura yellow, legs yellow-red, darkened to- 85 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 alia da eel aidan ai | > 2888 em s Figs. 30-38. Haliplus astrakhanus, holotype. - 30, dorsal view; 31, antenna; 32, elytral punctuation; 33, dorsal side of hind tibia; 34, prosternal process; 35, lateral view of prosternal process; 36, left paramere; 37, penis; 38, right paramere. 86 wards the coxae. Prosternal process flat, moder- ately strongly punctured, slightly narrowed near the coxae (fig. 34), lateral plicae complete (fig. 35). Metasternal process with a pit in the middle, weakly punctured (fig. 34). Setiferous striole '/, X the tibia length, with about 6 weak punctures, longer of the two tibial spurs 24 X the length of the first tarsal segment (fig. 33). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and paramere as in figs. 36-38, left paramere with a small solid parallel digitus, top of penis narrow and a little pointed. Etymology. - Named after the type locality. Distribution (fig. 304) Only known from the type-region Astrakhan in the south-west of the USSR. 5. Haliplus basinotatus Zimmermann (figs. 39-47) Haliplus basinotatus Zimmermann, 1924: 137. Lectotype & (here designated), [USSR] ‘Nikolsk Ussurijsk, Us- suri geb. Mandl, R. Mouchamps det. 62, Haliplus ba- sinotatus Zimm., Cotype’ (ISNB) [examined]. Haliplus basinotatus ssp. latiusculus Nakane, 1985a: 63. Holotype @, Yoshii, Fukuoka Pref., Kyushu, Japan, 10.vii.1957, N. Gyotoku leg. [not examined]. Haliplus basinotatus; Zimmermann 1924: 137, Zaitsev 1953: 75, Nakane 1985a: 63. Haliplus basinotatus ssp. latiusculus, Nakane 1987: 30. Diagnosis This species can be distinguished from related species by the dark basal band on the elytra and its large size. The Japanese subspecies latiusculus should differ from the continental form by having ‘the front margin of pronotum not or scarcely rounded-produced forwards and the punctuation and elytral pattern appearing somewhat different’ (Nakane 1985a). Description Length 3.5-4.0 mm, width 1.9-2.1 mm. Body widest just behind the well pronounced shoulders (fig. 39). Head. - Yellow to yellow-red, vertex usually dar- kened and nearly unpunctured, frons punctured. Distance between the eyes 1.3-1.4 X eye-width. Antennae (fig. 40) and palpi yellow-brown. Pronotum. - Yellow to yellow-red, front margin somewhat darkened or with a small brown blotch. Lateral borders slightly concave to slightly convex with very fine margins. In the anterior part with strong and dense punctures, basal punctures not wider than elytral punctures (fig. 39). VAN VONDEL: Palaearctic Liaphlus Elytra. — Yellow to yellow-red with distinct brown to nearly black markings on the intervals, along the suture, on the apex and along the base until the sixth puncture-row. The median spots are connected to the suture, but other spots can also show connections (fig. 39). Primary punctures strong, 28-31 punctures in the first row. Secondary punctures fairly strong. All punctures darkened (fig. 41). Ventral side. - Body yellow to yellow-red, legs yellow to yellow-red, darkened towards the brown trochanteres and coxae. Prosternal process flat, strongly and densely punctured and narrowed near the coxae (fig. 43), laterally with complete plicae (fig. 44). Metasternal process flat, in the middle behind the coxae with a shallow pit (fig. 43), mod- erately punctured. Setiferous striole on dorsal face of hind tibia with 11-20 punctures, sometimes in a strong groove making the separate punctures hardly recognizable, longer of the two tibial spurs about 24 X the length of the first tarsal segment (fig. 42). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 45-47. Distribution (fig. 311) Easternmost parts of the USSR (near Vladivos- tok), North Korea, Japan (ssp. latiusculus, accord- ing to Nakane 1985). Material examined. - 5 ex.- USSR: 1 @, 1 Q (lectotype and paralectotype) (ISNB). - North Korea: 29, 29.vii.1956, M. Magyar, Ch'ongjin (Tshondtsin) (ISNB); 1 ex., Tumen-ula, Russia/Korea border, 14.vii.1913 (ZMUC). 6. Haliplus chinensis Falkenstròm (figs. 48-56) Haliplus chinensis Falkenstròm, 1932: 191. Holotype 9, ‘China, N. O. Sichuan, Sven Hedins Exp. Ctr. Asien, Dr. Hummel, 22/5, Typus, Hal. chinensis n. sp. Typ. det. Falkenström’ (NHRS) [examined]. Haliplus chinensis; Falkenstròm 1933: 4 (again described as n. sp.), Zaitsev 1953: 71 (as ovalis) and 76 (as chinensis). Diagnosis This species is closely related to ovalis. It seems that ovalis is restricted to Japan, while the contin- ental specimens, usually considered to belong to ovalis, seem to be more related to chinensis, al- though only the female holotype of the latter spe- cies has the prosternal process slightly impressed. Despite the fact that all ovalis-like specimens that 87 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 | | I | @: ee 46 eae e a 88 re Figs. 39-47. Haliplus basinotatus, lectotype. - 39, dorsal view; 40, antenna; 41, elytral punctuation; 42, dorsal side of hind tibia; 43, prosternal process; 44, lateral view of prosternal process; 45, left paramere; 46, penis; 47, right paramere. 88 I have seen have a flat prosternal process I think that the continental specimens belong to chinensis. The division between the two is mainly based on the smoothly curved penis in the continental spe- cimens, while the specimens from Japan have the penis abruptly bent in the apical part. Description Length 3.9-4.2 mm, width 2.1-2.2 mm. Body oval, widest in the middle (fig. 48). Head. - Yellow to yellow-red, dark marking on vertex, weakly punctured, on the vertex with strong punctures. Distance between the eyes 1.4- 1.5 X eye-width. Antennae (fig. 49) and palpi yel- low-red. Pronotum. - Yellow, lateral borders straight, finely bordered. Densely, on the disc sparsely punc- tured, basal punctures widened and darkened, sometimes only slightly. Elytra. - Yellow. Suture, apical point and 9 or 10 blotches dark, central blotch on first and third in- terval sometimes connected to the suture (fig. 48). Primary punctures moderately strong, about 32-36 punctures in the first row. Secondary punctures fairly strong, apically almost as strong as the prim- ary punctures. All punctures darkened (fig. 50). Ventral side. - Body yellow to yellow-red, elytral epipleura yellow, legs yellow-red, slightly darkened towards the coxae. Prosternal process flat to clearly depressed in the middle of the apical part, strongly and densely punctured, narrowed near the coxae (fig. 52), lateral plicae complete (fig. 53). Metaster- nal process flat with a pit in the middle, rather weakly punctured (fig. 52). Setiferous striole on dorsal face of hind tibia about half of the tibia length, consisting of about 14 punctures, longer of the two tibial spurs % X the length of the first tarsal segment (fig. 51). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 54-56. Distribution (fig. 309) China, provinces of Sichuan, Shanxi, Jiangsu, Yunnan. Remarks Sato (1984) mentioned a subsp. sinicus Balfour- + Browne, but the description is unknown to me. Material examined. — 18 ex.- China: 1 9, Holotype 9. (NHRS); 1 8, 1 9, Nanjing, v.1928, J. G. Needham (ISNB); 1 6, 1 9, (Kiangau), C. E Wu (OLML); 1 6, 1 9, Lex. Shanghai; 1 ex., Yunnan Sen, Excoffier, 1898 (MNHN); 4 8, Kelan Pt. (1 ex. labelled as type of subsp. sinicus Balfour-Browne) (BMNH, MNHN); 5 ex., Xinjiang (Chin- kiang) (SMFD). VAN VONDEL: Palaearctic Liaphlus 7. Haliplus dalmatinus Miller (figs. 57-69) Haliplus dalmatinus Müller, 1900: 113. Lectotype & (here designated), [ Yugoslavia, Knin] ‘Kistanje, Müller D., Museo Civico di Trieste’ (MSNT) [examined]. Haliplus dalmatinus var. Weberi Miiller, 1900: 114. Lec- totype @ (here designated), [Yugoslavia] 'Mostarsko blato, TYPUS, Museo Civico di Trieste’ (MSNT) [examined ]. Haliplus dalmatinus, Zimmermann 1920: 307, 1924: 37, Bertrand 1928: 87, Scholz 1929: 16, Guignot 1933: 241, Burmeister 1939: 212, Zaitsev 1953: 71, Gueor- guiev 1958: 44, Franciscolo 1972: 63, Ienistea 1978: 294. Haliplus dalmatinus var. Webert, Zimmermann 1920: 308, 1924: 37, Bertrand 1928: 87, Scholz 1929: 16, Guignot 1933: 241, 1947: 51. Diagnosis This species is closely related to fulvus, but the latter has shorter metatarsi, a more narrowed pros- ternal process, in the males a longer digitus on the right paramere and generally a larger size. Description Length 3.2-3.9 mm, width 1.8-2.2 mm. Body oval, sides somewhat parallel, widest in the middle (fig. 57). Head. - Yellow-red with dark vertex, weakly and sparsely, behind the eyes stronger and more den- sely punctured. Distance between the eyes 1.4-1.7 X eye-width. Antennae (fig. 58) and palpi yellow- red. Pronotum. - Yellow, lateral borders straight to slightly convex, finely bordered, except in the mid- dle fairly strongly punctured, basally with a row of widened punctures, basal and discal punctures dar- kened. Elytra. - Yellow with variable pattern of brown blotches in the even intervals, sometimes con- nected to each other, most of the suture and apex darkened (figs. 57, 66-69). Fairly strong primary puncture-rows, about 30 punctures in the first row. Secondary punctures relatively strong. All punc- tures darkened (fig. 59). Ventral side. - Body yellow-red, elytral epipleura yellow, legs yellow-red to brown towards the coxae. Prosternal process flat, rather strongly punctured, narrowed near the coxae (fig. 61), lateral plicae complete (fig. 62). Metasternal process flat or slightly grooved, shallow pit in the middle, sparsely punctured (fig. 61). Setiferous striole on dorsal face of hind tibia with 4-7 punctures, longer of the two tibial spurs % to 24 X the length of the first tarsal segment (fig. 60). Length of hind tarsus 1.3-1.5 X the tibia length, first tarsal segment 1.6-1.7 X the length of the second segment. 89 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 48-56. Haliplus chinensis (48-53, holotype; 54-56, from Nanjing). - 48, dorsal view; 49, antenna; 50, elytral punctuation; 51, dorsal side of hind tibia; 52, prosternal process; 53, lateral view of prosternal process; 54, left paramere; 55, penis; 56, right paramere. 90 VAN VONDEL: Palaearctic Liaphlus UU GOA 159 Figs. 57-65. Haliplus dalmatinus, lectotype. - 57, dorsal view; 58, antenna; 59, elytral punctuation; 60, dorsal side of hind tibia; 61, prosternal process; 62, lateral view of prosternal process; 63, left paramere; 64, penis; 65, right paramere. 91 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 e ee oe DOS 5 29 009 09 naer ec 29 0 a PI S AUS SI SIAE SR “ di à “ i] 1 DE 5 Oe 0 1 i + O: Kl DEE 4 "I 7 whe 3 3 000 d bi i u 12] Soi Uh 5 E . È dia tive $ Al an x . ESP VETTEN ety ee TOR! VENA co sg ave ga labs hole LAT Figs. 66-69. Haliplus dalmatinus, elytra. - 66, from Kistanje; 67, from Zara; 68, from Knin; 69, from Skutari. Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 63-65, length of distal process of right para- mere |. X the length of the basal part (fig. 65). Distribution (fig. 301) Along the coast of Yugoslavia and Albania, on the island Corfu (Greece), Turkey, Italy? (identifi- cation not sure). Remarks Since the description of this species various au- thors have regarded it as a variety of fulvus. Some authors considered dalmatinus a variety or subspe- cies of variegatus. Although in some cases dalma- tinus is very hard to distinguish from small spe- cimens of fulvus I consider dalmatinus a separate species and I regard the var. weberi as an occasional aberration. The elytral maculation is very variable and ranges from almost absent to very extended. One of the syntypes of Haliplus zimmermanni from Italy might belong to this species. Specimens from Zadar (Zara), labelled as type, are not considered to belong to the syntype-series as this locality is not mentioned in the original des- cription. Material examined. - 36 ex. - Yugoslavia: 34, 29 (lec- totype and paralectotypes); 3 ex., surr. of Zadar (Zara), Müller, (1 labelled as Typus); 2 ex., Krk (Veglia), Neto- 92 litzky; 1 ex., Dalmatia, Rab (Arbe), 20.ix.1910 (MSNT); 1 ex. (paralectotype), Knin (Kistanje), Miiller D., Dal- matin det. J. Müller m, Cotypus (ZSMC); 1 & (paralecto- type), Knin D., Müller, Cotypus; 1 ex., surr. Zadar (Zara), Müller, Cotypus; 1 6, Dalmatia; 1 @, Dalmatia, Omis (Almissa), v.1911, Schatzm.; 3 ex. (lectotype and paralec- totypes of dalmatinus var. weberi), Mostarsko blato, Ty- pus, Haliplus v. Weberi (MSNT); 1 ex., Sinj, Dalmatia (NHMW). — Albania: 2 4, Shkodér (Skutari), I. See-Ufer; 2 &, Vraka near Shkodër (NHMW). - Greece: 1 ex., Kérkira (Corfu), J. Sahlb., 1035, syntype Haliplus leopardinus Sahlberg (1900) (zsMc). - Turkey: 3 6, 2 9, Yüksekova, 3.vi.1987, M. Jäch; 5 6, Kastamonu, 25.vi.1989, M. Jäch (NHMW). — Italy : 1 9 (syntype of Haliplus zimmermanni Gschwendtner, 1921), Salo, 23.v.1909, Type 1921, det. L. Gschw., Type, Haliplus ?dalmatinus Müller det. B. J. v. Vondel 1987 (ZSMC). 8. Haliplus davidi sp. n. (fig. 70-78) Type material: Holotype &, [China, Beijing] ‘Nord Pekin, A. David, 1865, Museum Paris, Haliplus Sharpi Westw., M. Régimbart det. 1898, 227.65 [round label] (MNHN); Paratype 9, China, Yunnan (MNHN); Paratype 8, Beijing (Peping), CF. Wu (OLML); Paratype 4, Hei- longjiang (Mandsch.), Harbin, 28.ix.1930, v. Jettmer (ZMHB). Diagnosis Male specimens can be distinguished from di- ruptus by the presence of a digitus on the left paramere. I am not always able to separate the females of these species. VAN VONDEL: Palaearctic Liaphlus Figs. 70-78. Haliplus davidi, holotype. - 70, dorsal view; 71, elytral punctuation; 72, dorsal side of hind tibia; 73, prosternal process; 74, lateral view of prosternal process; 75, hind coxal plate; 76, left paramere; 77, penis; 78, right paramere. 93 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 79-87. Haliplus diruptus, holotype. - 79, dorsal view; 80, antenna; 81, elytral punctuation; 82, dorsal side of hind tibia; 83, prosternal process; 84, lateral view of prosternal process; 85, left paramere; 86, penis; 87, right paramere. 94 Description Length 3.2 mm, width 1.7 mm. Body oval, widest in the middle (fig. 70). Head. - Yellow-brown, brown blotch on vertex, densely punctured, unpunctured band between the hind parts of the eyes. Distance between the eyes 1.3 X eye-width. Antennae yellow (missing in ho- lotype), palpi yellow-brown. Pronotum. - Yellow-brown. Lateral borders straight, clearly bordered. Width at the base 2.0 X the length in the middle and 1.6 X the width in front. Weakly and densely, along the base more strongly punctured, some basal punctures dar- kened. Beside the middle of the base slightly im- pressed (fig. 70). Elytra. - Yellow-brown, with darkened suture and apex and several blotches (fig. 70). Primary puncture-rows moderately strong, wider near the base, basal punctures in the second to fourth row widened, about 35 punctures in the first row. Se- condary punctures moderately strong, in some in- tervals very sparse, accompanied by very small punctures (fig. 71), all punctures darkened. Ventral side. - Yellow-red to yellow-brown, ely- tral epipleura yellow, legs yellow-brown, some dar- kening towards the coxae. Prosternal process at the apex strongly impressed, coarsely punctured, pos- teriorly little narrower than anteriorly, slightly narrowed near the coxae (fig. 73), lateral plicae complete (fig. 74). Metasternal process grooved and strongly impressed in the middle, rather strongly punctured (fig. 73). Coxal plates strongly and densely punctured, small punctures between the normal ones, which are provided with hairs, along the suture with a weakly punctured area, apical parts of the plates dilated (fig. 75). Setiferous striole 4 of the tibia length, with about 5 punc- tures, longer of the two tibial spurs % the length of the first tarsal segment (fig. 72). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 76-78, left paramere with a short solid digitus. Etymology. — Named after the collector A. David. Distribution (fig. 312) Only known from China: Yunnan, Beijing and Harbin. 9. Haliplus diruptus Balfour-Browne (figs. 79-90) [Haliplus simplex sensu Régimbart 1899, nec Clark 1863. Misidentification]. Haliplus diruptus Balfour-Browne, 1946: 436. Holotype VAN VONDEL: Palaearctic Liaphlus 8 ‘China, Tientsin’ [Tianjin] ‘EF. M. Thomson, 1904- 229, Haliplus simplex Rég.-Zimm., Haliplus diruptus nom. nov. J. Balfour-Browne det., Type’ (BMNH) [examined]. Haliplus simplex; Zimmermann 1924: 136, Takizawa 1931: 142, Zaitsev 1953: 74, Sato 1960: 252, 1984: 2, Cho 1969: 167, Yoon 1988: 621. Haliplus diruptus; Guignot 1954: 563. Diagnosis A very variable species. Specimens with and without black blotches or bands on the base of the elytra and specimens with weak or with very strong and close punctures on the head are considered to belong to this species. Specimens with a very re- duced and weakened maculation resemble davidi, from which males can be distinguished by the lack of a small solid digitus on the left paramere, but distinguishing females can be problematic. Description Length 2.8-3.5 mm, width 1.6-2.0 mm. Body oval, widest in the middle (fig. 79). Head. - Yellow to yellow-brown, vertex slightly darkened, puncturation weak and fairly sparse to strong and dense. Distance between the eyes 1.3- 1.5 X eye-width. Antennae (fig. 80) and palpi yel- low-red. Pronotum. - Yellow to yellow-red. Lateral borders straight to slightly convex, finely bordered. Punctures moderately strong to strong along the base and in the anterior central part, basal and anterior punctures darkened, remaining punctures sometimes darkened. Elytra. - Yellow to yellow-brown, variable mac- ulation: darkening along suture and on apex, a central blotch on the suture and small blotches on the even intervals; inextreme forms the maculation can be very extended and includes a large blotch or even a complete band along the base (figs. 88-90). Primary punctures moderately strong, first row slightly weaker than second to fourth or even fifth row, 30-40 punctures in the first row. Secondary punctures moderately strong, rows consisting of normal and very small ones, sometimes very strong along suture. All punctures darkened (fig. 81). Ventral side. - Body yellow to yellow-brown, elytral epipleura yellow, legs yellow to yellow- brown, slightly darkened near the coxae. Proster- nal process clearly impressed in the apical part, strongly narrowed near the front coxae (fig. 83), lateral plicae complete (fig. 84), fairly strongly punctured. Metasternal process flat with a pit in the middle, sparsely punctured (fig. 83). Setiferous striole on dorsal face of hind tibia about % X the tibia length, containing about 8 punctures, longer of the two tibial spurs about % X the length of the first tarsal segment (fig. 82). 95 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 88-90. Haliplus di- ruptus, elytra. - 88, from Beijing; 89,90, from Ton- kin. oo OC SO OG oO pons Os ORDO sy Ce eR OO Se . ER. «* Se Os Male sexual characters. — First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 85-87. Distribution (fig. 311) Easternmost part of China, provinces of Hebei, Heilongjiang, Hunan and Jiangsu. Extreme south- eastern part of the USSR: Maritime Territory. Vietnam. Guignot (1954) gives Burma, Shan State. Material examined. — 81 ex. - China: 1 &, Holotype; 1 Q (paratype), Tianjin (Tientsin), vi.1902, F. M. Thom- son; 1 6, Harbin, 25.viii.1952 (BMNH); 1 Ó, 1 9, Beijing (Peping), C. E Wu (oLML); 1 6, 1 9, Tianjin (Tientsin) (SMED); 1 ex., Tianjin (Tien-Tsin); 1 ex., Beijing (Nord Pekin), A. David 1865; 4 ex., Shanghai; 1 ex., Hengshan (Hontsang), Weber; 1 @, (Zi-ka-wei) (MNHN); 1 ex, Yangzhou (Yang Chow Ku), 20.vi.1947, Sunabricotier (ISNB); 1 &, Beijing (Peping), C. F Wu (OLML). - Korea: 1 ex., 11.vii.1956, M. Magyar, Sariwon, Type, Haliplus sharpi var. homologus n. var. cf. texte, R. Mouchamps det. 63 [unpublished name] (ISNB). — Vietnam: 62 ex., Tonkin, Hoa Binh region, vii.1940 (part of material), A. de Cooman (ISNB, MNHN). — Without locality: 1 ex. (MNHN). 10. Haliplus excoffieri sp. n. (fig. 91-101) Type material: Holotype g, [China] ‘Yunnan-Sen, Mgr Excoffier, 1898, Dr. Régimbt vidit 1898, Museum Paris ex coll. Oberthiir’. Paratypes: 1 6,3 9, same data (MNHN). Diagnosis This species is closely related to chimensis and ovalis, but the distance between the eyes is usually 96 less than 1.4 X eye-width, the darkened suture is only reaching to the first secondary puncture-row and in the males the penis is clearly widened before the apex. Description Length 3.7-3.8 mm, width 1.9-2.0 mm. Body oblong oval, tapering apically, widest in or just before the middle (fig. 91). Head. - Brown, dark marking on vertex, rather densely punctured, unpunctured band between the hind parts of the eyes. Distance between the eyes 1.2-1.4 X eye-width. Antennae yellow to yellow- brown (fig. 92), palpi yellow-brown. Pronotum. - Yellow to yellow-brown, lateral borders straight, clearly bordered. Width at the base about 2.0 X the length in the middle and 1.6 X the width in front. Sparsely, anteriorly densely, basally strongly punctured, on the disc almost un- punctured. About ten of the darkened basal punc- tures widened, sometimes slightly impressed along the central part of the base (fig. 91). Elytra. - Yellow to yellow-brown, suture dar- kened until the secondary puncture-row, dark blotches on the apex and in mainly the even inter- vals. Primary puncture-rows moderately strong, second to fourth puncture-row getting stronger basally, 32-36 punctures in the first row. Secondary puncture-row along the suture rather weak, but very dense in the apical half, on the other intervals sometimes relatively strong (fig. 93). All punctures darkened. Ventral side. - Yellow to yellow-red, elytral epi- pleura yellow, legs yellow-brown, darkened to- wards the coxae. Prosternal process slightly con- vex, strongly narrowed near the coxae, densely punctured (fig. 96), lateral plicae complete (fig. 97). Metasternal process flat, strong impression in the middle, weakly punctured (fig. 96). Coxal plates strongly punctured, along the suture weakly punc- tured (fig. 98). Hind margins of fifth and sixth abdominal segment with complete double punc- ture-rows (fig. 98). Setiferous striole almost half of the tibia length, with about 10 groove-like con- nected punctures, longer of the two tibial spurs % the length of the first tarsal segment, tibia 3 X the length of the first tarsal segment (fig. 95). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on the ventral side. Penis and parameres as in figs. 99- 101. Etymology. — This species is named after the collector Mgr J. C. Excoffier. Distribution (fig. 312) Only known from China, Yunnan-province. 11. Haliplus eximis Clark (figs. 102-110) Haliplus eximis Clark, 1863: 418. Syntypes, Korea, Clark (depository unknown; probably in SAMA or MAMU). Haliplus modestus Zimmermann, 1924: 139. Holotype 6, ‘Sumatra, Type, Holotype Staatssamml. Miinchen’ (ZSMC) [examined]. Syn. n. Haliplus hiogoensis Kano & Kamiya, 1931: 1. Types burned during World War II (Dr. T. Nakane, pers. comm.) [not examined]. Haliplus emmerichi Falkenstròm, 1936: 79. Holotype 9, [Kangdin, Sichuan] ‘Nitou Tatsienlu, Szechuan China , Typus, Haliplus Emmerichi Falk. 9, det. Falken- ström’ (NHRS) [examined]. [Haliplus flavicollis; auctt., nec Sturm 1834: Wu 193:342. Misidentification |. Haliplus eximis; Zimmermann 1920: 305, 1924: 132, Takizawa 1931: 140, Guignot 1950: 97, Zaitsev 1953: 77, Cho 1969: 166, Sato 1984: 2, Nakane 1985a: 63, 1985b: 40, 1987: 29, Yoon 1988: 623. Haliplus modestus, Zimmermann 1927: 2. Haliplus hiogoensis; Sato 1984: 2, Nakane 1985b: 40. Haliplus emmericht, Guignot 1950: 97, Zaitsev 1953: 69, Sato 1984: 2. Diagnosis This species can be distinguished from related species in the East Palaearctic and Oriental region by the total lack of maculation between the elytral puncture-rows. It resembles the West Palaearctic flavicollis, which however lacks the lateral plicae in the anterior part of the prosternal process. Description Length 3.4-3.9 mm, width 1.9-2.1 mm. Body oval, widest in or just before the middle (fig. 102). Head. - Yellow-brown to rust-coloured, mode- VAN VONDEL: Palaearctic Liaphlus rate weakly punctured. Distance between the eyes 1.3-1.4 X eye-width. Antennae (fig. 103) and palpi yellow-brown. Pronotum. - Yellow to rust-coloured. Lateral borders straight to slightly convex, bordered, ba- sally wider than base of elytra, strongly tapering anteriorly. Width 2.1-2.2 X the length in the mid- dle. With strong punctures except on the unpunc- tured disc, basal punctures partly widened, basal and anterior punctures darkened. Elytra. - Yellow to rust-coloured. Strong primary puncture-rows, about 32 punctures in the first row. Secondary punctures relatively strong, as dense and apically almost as strong as the primary punctures (fig. 104). Except for the partly darkened suture, no markings present. Ventral side. - Yellow, sternites yellow-brown, legs yellow-brown to rust-coloured. Prosternal process broad and flat, slightly narrowed near the coxae, strongly punctured (fig. 106), lateral plicae complete (fig. 107). Metasternal process flat with a pit in the middle, sparsely punctured (fig. 106). Setiferous striole on dorsal face of hind tibia with about 12 punctures, longer of the two tibial spurs % X the length of the first tarsal segment (fig. 105). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 108-110. Distribution (fig. 308) China (prov. Sichuan, Xinjiang, Zhejiang, Zait- sev (1953) gives Shanghai), Korea, Japan (Hon- shu, Kyushu), Vietnam, Sumatra. Remarks The specimen from China, present in BMNH and labelled as Type of eximis, does not belong to Clark’s two syntypes, described from Korea. The holotype of modestus, described from Sumatra is here considered conspecific with exzmzs. Material examined. - 16 ex. - China: 1 4, Type (BMNH); 1 ex., Kiangau, C. FE Wu (OLML); 1 9, Holotype Haliplus emmerichi (NHRS); 4 ex., North-west. China, Xinjiang (Chinkiang), Cotypus (SMFD); 1 6, Zhejiang, Zhoushan (Chekiang, Chusan), 15.iv.1931 (MNHN). — Ja- pan: 10 ex., Aobadai, Aira I, Kagoshima Pref., 26.ix.1985. E. Matsui (Vondel). - Korea: 1 ex., Tshou-Bon San, 1.vii.1956, M. Magyar (ISNB). - Vietnam: 6 ex., Tam Dao, Tonkin, H.Perrot; 1 ex., Tonkin, Hoa Binh, iii.1940, A. de Cooman (ISNB). - Indonesia: 1 9, holotype of modestus (ZSMC). 12. Haliplus flavicollis Sturm (figs. 111-119) 97 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ce eo oe °° = O Gy: Ma 0 9 a see 8 e © 0 o o punctuation; 94, fore claws; 95, prosternal process; 98, hind coxal plate and sternites; 91, dorsal view; 92, antenna; 93, elytral ; 97, lateral view of ; 96, prosternal process 98 ; 100, penis; 101, right paramere. Figs. 91-101. Haliplus excoffieri, holotype. - dorsal side of hind tibia 99, left paramere VAN VONDEL: Palaearctic Liaphlus 103 POCO d Ki À ©: CR OM SEE, II AR Ted Er GEO Eis © eovoe oe À OTO 0- OOO DO ehehe SE 2 3,0 © 0.0 0 KO 10900 OOO OO mc: e e ® LOS 28 8 6 00000600 5, DO ORE ° soe 26360 0.007 Sa 4 5 edad OAT . = e OTO e. ° © © na o DA © 600009099. EVO Lee OMC frere er Vo. © oie . , elytral punctuation; 105, dorsal side of ; 108, left paramere; 109, penis; 110, right 110. Haliplus eximis, China. - 102, dorsal view; 103, antenna; 104 ; 106, prosternal process; 107, lateral view of prosternal process Figs. 102- hind tibia paramere. 99 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 111-119. Haliplus flavicollis, Hendrik Ido Ambacht, Netherlands. - 111, dorsal view; 112, antenna; 113, elytral punctuation; 114, dorsal side of hind tibia; 115, prosternal process; 116, lateral view of prosternal process; 117, left paramere; 118, penis; 119, right paramere. 100 Haliplus flavicollis Sturm, 1834: 150. Types not located, probably lost as most of the Sturm-types [not exam- ined]. (Haliplus ferrugineus, sensu auctt., nec Linnaeus, 1767. Misidentification |. (Haliplus impressus, sensu auctt., nec Fabricius,1787. Misidentification]. Haliplus flavicollis var. pallida Semenov, 1904: 202. Types probably in ZMAS [not examined]. Haliplus flavicollis var. expallidus Zaitsev, 1907: 122 (Replacement name for pallida Semenov). Haliplus unicolor Fiori, 1914: 188. Types not located [not examined |. Haliplus flavicollis, Aubé 1836: 24, 1838: 12, Schaum 1848: 42, Thomson 1854: 183, Seidlitz 1887: 31, Miiller 1900: 115, Apfelbeck 1904: 361, Fiori 1904: 196, Zimmermann 1920: 305, 1924: 132, Guignot 1928: 143, 1933: 230, 1947: 50, Kinel 1929: 219, Scholz 1929: 15, Kolossov 1931: 116 (var. russkyi Stange, nom. nud.), Balfour-Browne 1936: 75, 1938: 12, 1940: 148, 1951: 1, 1953: 5, FE Burmeister 1939: 212, E. Burmeister 1982: 94, 1987: 184, Csiki 1946; 559, Zaitsev 1953: 68, Gueorguiev 1960: 23, Lindroth 1960: 42, Freude 1971: 15, Seeger 1971a: 409, 1971b: 532, 197 1c: 165, Franciscolo 1972: 61, 1979: 117; Jack- son 1973: 253, Ienistea 1974: 208, 1978: 294, Mino- ransky & Dzumailo 1974: 26, Galewski 1976: 31, Konev 1976: 56, Mateleshko 1977: 68, Fichtner 1981: 327, Holmen 1981: 2, 1987: 114, Leblanc 1984: 8, Lundberg 1986: 11, Zakharenko & Moroz 1988: 283, Belyashevskiy 1989: 124, Vondel 1989a: 58, 1989b: 17. Haliplus flavicollis var. expallidus; Zimmermann 1920: 305, Guignot 1933: 239, Zaitsev 1953: 68. Haliplus unicolor; Burmeister 1939: 214. Diagnosis This species is related to eximis and rubidus. From eximzs it can be distinguished by the lack of lateral plicae in the anterior part of the prosternal process. In rubidus the basal elytral punctures are larger than the distance between the punctures of the first and second row in the basal part, while in flavicollis these punctures are clearly smaller than that distance. In specimens from Morocco, how- ever, that difference is less clear. The males of flavicollis have the left paramere narrower at the base than in rubidus. Specimens with vague blotches on the elytra can be distinguished from fulvus by the lack of the lateral plicae in the anterior part of the prosternal process. Description Length 3.6-4.1 mm, width 1.8-2.1 mm. Body fairly slender, sometimes subparallel, widest before the middle (fig. 111). Head. - Yellow to rust-coloured, sparsely punc- tured. Distance between the eyes 1.4-1.6 X eye- width. Antennae (fig. 112) and palpi yellow to yellow-red. Pronotum. — Yellow to rust-coloured. Lateral VAN VONDEL: Palaearctic Liaphlus borders straight, finely bordered. Sparsely, ante- riorly and basally more densely punctured, basal irregular puncture-row about as strong as basal elytral punctures. All punctures usually black (fig. bly Elytra. - Yellow to rust-coloured, strong primary puncture-rows, about 32 punctures in the first row. Secondary rows with relatively strong and between them very weak punctures (fig. 113). All punctures black. Suture weakly darkened, no clear markings on the intervals, sometimes vague blotches on the apical half. Ventral side. - Body yellow to yellow-brown, elytral epipleura yellow, legs yellow-brown, femora darkened basally, trochanteres dark. Prosternal process flat, wide apically, strongly narrowed near the coxae (fig. 115), lateral border without plicae before the coxae (fig. 116), strongly and densely punctured. Metasternal process flat with a strong impression in the middle, sparsely punctured (fig. 115). Setiferous striole on dorsal face of hind tibia long and grooved, separate punctures hardly recog- nizable, longer of the two tibial spurs about % X the length of the first tarsal segment (fig. 114). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs hardly widened later- ally, slightly widened ventrally, scaly hairs on ven- tral side. Penis and parameres as in figs. 117-119. Female: Elytra completely and very densely co- vered with micropunctures. Biology In clean and clear stagnant or very slowly run- ning water of lakes, ditches, canals and pools with rich vegetation, especially Characeae (Seeger 1971c), Najadaceae (Burmeister 1939) or Ca- bomba-beds in Kazakhstan (Konev 1976). Also recorded from brackish water. The adults feed on eggs and larvae of Chironomidae (Seeger 1971c), aquatic macro-invertebrates as Asellus (only when dead or dying), Stalis larvae (probably predated), dead waterbeetles, even of their own species and probably Characeae or filamentous algae. Seeger (1971a, b, c) provided much information on the life-cycle. The egg has been described by Seeger (1971a), the 2nd and 3rd-stage larvae and the pupa have been described by Bertrand (1928). This spe- cies is able to fly (Jackson 1973, taken at light). Distribution (fig. 295) All Europe except the north of Norway, the utmost north of Sweden, the north of Finland, Iceland, Spain and Portugal. Turkey. USSR: Euro- pean part, Western Siberia except the extreme north, Transcaucasia, Kazakhstan, China (Guignot 1947), Mongolia. Records from Morocco probably concern specimens of rzbidus, which is not always 101 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 e . © o ©0006" ’ Austria. - 120, dorsal view; 121, antenna; 122, elytral punctuation; 123, dorsal ; 125, lateral view of prosternal process Figs. 120-128. Haliplus fulvus, Gars, : 126, left paramere; 127, penis; 128, side of hind tibia; 124, prosternal process right paramere. 102 easy to distinguish from flavicollis. Egypt? (Hol- men 1987). Ali (1976) records specimens of fulv1- collis from Iraq, but according to the characters he used in his identification-key he obviously meant flavicollis. Material examined. — 881 ex. — Austria: 97 ex. (CNCI, NHMW, ZMAN, RMNH). — Belgium: 7 ex. (RIN, RMNH, Jon- gema). — Czechoslovakia: 20 ex. (NHMW, RMNH). — France: 23 ex. (CNCI, NHMW, RIN, RMNH, Jongema, Von- del). - Germany: 24 ex. (NHMW, ZMAN, RMNH). — Great Britain: 9 ex., (CNCI, BMNH, ZMAN, Vondel). — Greece: 2 ex. (ZMAN). — Hungary: 2 ex. (CNCI). - Ireland: 1 ex. (BMNH). — Italy: 4 ex. (CNCI, Jongema, RMNH, NHMW). — Luxembourg: 1 ex. (ZMAN). - Mongolia: 1 &, Uiir Gol, 28.ix.1897, Frenzel (ZMHB). — Netherlands: 649 ex. (ZMAN, MOG, FNML, NHMR, RIN, RMNH, Berge Hene- gouwen, Cuppen, Drost, Hielkema, Huybregts, Jongema, Onder de Linden, Schilthuizen, Vallenduuk, Vondel). — Poland: 13 ex. (NHMW, RMNH).- Rumania: 1 ex. (RMNH). - Switzerland: 8 ex. (ZMAN, Rotzer). - USSR: 1 ex., Panfilov, Kazakhstan; 1 ex., Kiyev, Ukraine, 27.ix.1988, M. Jäch (NHMw).- Locality not traced or unknown: 17 ex. (BMNH, NHMW, RMNH). 13. Haliplus fulvus (Fabricius) (figs. 120-131) Dytiscus fulvus Fabricius, 1801: 271. Types not located, probably lost (Zimsen 1964: 70) [not examined]. Dytiscus interpunctatus Marsham, 1802: 429. Lectotype Q (here designated), interpunctatus (BMNH) [examined]. Haliplus lapponum Thomson, 1854: 184. Lectotype (here designated), H. ferrugineus, coll. Lund, H.ferrugineus Zett., ?syntype (MZLU, Zetterstedt-collection) [examined |. Haliplus pyraeneus Delarouzée, 1857: xcv. Types from Lac Noire near Cauterets (Hautes Pyrénées) are prob- ably in MNHN [not examined]. Material from NHMW, labelled as TYPUS and COTYPUS is collected in 1881 and thus can not belong to the syntype-series. Haliplus lapponum var. niger Seidlitz, 1887: 31. Types not located [not examined]. Haliplus fulvus var. carlittensis Régimbart, 1901: 323. Types probably in MNHN [not examined]. Haliplus subguttatus Crotch, 1873: 384. Syn. n. Haliplus fulvus a. multistriatus Scholz, 1915: 232. Types not located [not examined]. Haliplus fulvus Sparre-schneideri Munster, 1922: 15. Lectotype @ (here designated), [ Norway, Malselv] ‘Bjerkeng, 12.6.97, H. Schneider: i. lit fulv. Schneideri det. Ganglbauer, COTYPUS’ (NHMw) [examined]. Haliplus fulvus v. unicolor Munster, 1922: 14. Types not located [not examined]. Haliplus salinarius Wallis, 1933: 56. Holotype 6, [Canada] ‘Oyosoos, B.C., V-4-1929° (CNCI) [exami- ned]. Syn. n. [Haliplus ferrugineus; sensu auctt., nec Linnaeus, 1767. Misidentification]. Haliplus fulvus; Clairville 1806: 220, Sturm 1834: 148, Babington 1836: 177, Erichson 1837: 183, Thomson 1854: 183, Westhoff 1881: 43, Seidlitz 1887: 31, Ap- VAN VONDEL: Palaearctic Liaphlus felbeck 1904: 360, Escalera 1914: 57, Zimmermann 1920: 307, 1924: 134, Bertrand 1928: 87, 1949: 25, Guignot 1928: 144, 1933: 230, 1947: 51, Scholz 1929: 16, Balfour-Browne 1936: 74, 1938: 11, 1940: 146, 1951: 4, 1953: 5, Pearce 1939: 34, F. Burmeister 1939: 212, Horion 1941: 365, Brinck 1942: 5, Strand 1944: 122, Csiki 1946: 560, Jackson 1952: 66, 1956: 87, 1973: 253, Zaitsev 1953: 70, Riha 1955: 57, Gueorguiev 1958: 44, 1960: 23, 1981: 401, Lindroth 1960: 42, Lagar Mascaro 1968: 72, Freude 1971: 15, Seeger 1971b: 558, 1971c: 165, Ienistea 1974: 208, 1978: 294, Minoransky & Dzumailo 1974: 26, Alfieri 1976: 38, Galewski 1976: 29, Konev 1976: 56, Gislason 1977: 159, Mateleshko 1977: 68, Franciscolo 1979: 119, Fichtner 1981: 327, Holmen 1981: 2, 1987: 115, Leb- lanc 1984: 40, Lundberg 1986: 11, E. Burmeister 1987: 184, Zakharenko & Moroz 1988: 283, Belyashevskiy 1989: 124, Vondel 1989a: 58, 1989b: 17. Haliplus interpunctatus, Zimmermann 1920: 307. Haliplus lapponum; Seidlitz 1887: 31, Zimmermann 1920: 308, Bertrand 1928: 87, 1949: 25, Guignot 1928: 144, 1930: 73, 1933: 231, 1947: 51, Kinel 1929: 219, Scholz 1929: 16, Balfour-Browne 1936: 74 1938: 11, 1940: 147, Burmeister 1939: 213, Pearce 1939: 34, Horion 1941: 365, Brinck 1942: 5, Strand 1944: 122, Zaitsev 1953: 71, Riha 1957: 57, Lagar Mascaro 1968: 72, Ienistea 1978: 294, Lundberg 1986: 11. Haliplus pyraeneus, Marseul 1882: 106, Zimmermann 1920: 308, Bertrand 1928: 87, Kinel 1929: 219, Scholz 1929: 16, Burmeister 1939: 213. Haliplus subguttatus; Blatchley 1910: 203, Roberts 1913: 101, Zimmermann 1920: 139, Wallis 1933: 54, 1973: 100, J. Balfour-Browne 1948: 158, Hatch 1953: 192, Ruette 1970: 57, Hilsenhoff & Brigham 1978: 21, White et al. 1985: 357, Larson 1987: 107. Haliplus lapponum var. niger; Scholz 1929: 16, Guignot 1933: 242. Haliplus fulvus var. carlittensis, Zimmermann 1920: 308, Bertrand 1928: 87, Guignot 1928: 144, 1930: 73, 1933: 242, Kinel 1929: 219, Scholz 1929: 16, Burmeister 1939: 213, Pearce 1939: 34. Haliplus fulvus a. multistriatus, Zimmermann 1920: 308, Bertrand 1928: 87, Guignot 1933: 241. Haliplus fulvus sparre-schneideri, Guignot 1933: 241, Strand 1944: 122, Zaitsev 1953: 71, Döbler 1976: 182 (as schneideri), Ienistea 1978: 294. Haliplus fulvus v. unicolor; Falkenstròm 1941: 88. Haliplus salinarius; Hatch 1953: 192, Ruette 1970: 56, Larson 1987: 107. Haliplus ferrugineus; Gyllenhal 1808: 546, Zetterstedt 1828: 236, Stephens 1829: 40, Aubé 1836: 22, 1838: 10, Babington 1836: 176, Zimmermann 1920: 308. Diagnosis This species is very variable in size, maculation and coloration. Of all Haliplidae it has the largest distribution, covering most of the palaearctic and the north of the nearctic region. Mainly because of the variation in elytral maculation a number of varieties, subspecies and separate species such as lapponum (fig. 129) and pyraeneus (fig. 130) have been described. A range of intermediate forms be- tween the latter species and fulvus has been exam- ined and I can only confirm what other authors like 103 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ole sel She cheb? ERS ee e Figs. 129-131. Haliplus fulvus, elytra. - 129, lec- totype of H. lapponum, 130, from Cauterets, France, the type locality of H. pyraeneus; 131, lec- totype of H. fulvus sparre-schneideri. Balfour-Browne (1936) and Holmen (1987) al- ready concluded, that they belong to the same spe- cies. Description Length 3.6-4.5 mm, width 2.0-2.4 mm. Body long oval, widest in the middle (fig. 120). Head. - Yellow-red, brown or rust-coloured, puncturation weak to moderately strong, between the eyes sparser, behind the eyes stronger and denser punctured. Distance between the eyes 1.6- 1.9 X eye-width. Antennae (fig. 121) and palpi yellow. Pronotum. - Yellow to rust-coloured. Lateral borders straight to slightly convex, finely bordered. Punctures moderately strong, on the disc almost absent, the basal row widened and slightly to strongly darkened. Elytra. - Yellow to rust-coloured, very variable pattern of light to dark brown blotches in the even intervals, sometimes the blotches are hardly or not present or they are very strong and then covering nearly the entire elytra (described as sparre- schneideri) (fig. 131). Primary punctures moder- ately strong, sometimes stronger in the basal part. Secondary rows irregular and containing two types of punctures, the usual ones and very fine ones (fig. 122). All punctures darkened. Ventral side. - Body yellow, yellow-red or rust- 104 9.0 0.0 0.0 9.9 a è Bag geen coloured, elytral epipleura yellow, legs yellow to rust-coloured, darkened towards the coxae. Pros- ternal process flat to slightly convex, narrowed near the coxae (fig. 124), lateral plicae complete (fig. 125), with moderately strong to very strong and dense punctures. Metasternal process flat with a pit in the middle (fig. 124), with moderately strong and dense punctures. Setiferous striole on dorsal face of hind tibia grooved, stretched over about % of the length, consisting of 10 to 20 often hardly recognizable punctures, longer of the two tibial spurs % X the length of the first tarsal seg- ment (fig. 123). Length of hind tarsus, including claws, about 1.2 X the tibia length, first tarsal segment 1.5-1.9 X the length of the second seg- ment. Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 126-128, length of the distal process of the right paramere % X the length of the basal part (fig. 128). Biology In clean clear or peaty stagnant or slowly run- ning water of lakes, pools, canals, rivers and ditches with rich vegetation of especially Characeans. Even recorded from brackish water. The larvae probably feed on Characeans such as Nitella, while the adults are omnivorous with a diet of algae, pollen and invertebrates like Oligochaeta (Seeger 1971b). The third-stage larva was first described by Schigdte (1864). The beetle has been seen flying and has been caught at light. Distribution (fig. 292) All of Europe, including Iceland. USSR: Euro- pean part and Siberia as far as Yakutia in the east and Transcaucasia and Kazakhstan in the south. Turkey. North Africa: Morocco, Algeria. The re- cords of Syria (Burmeister 1939) and Iraq (Ali 1976) possibly concern maculatus, which is cer- tainly a different species. Canada (states of British Columbia, Alberta, North West Territories, Saskat- chewan, Manitoba, Ontario, Quebec, Nova Scotia and New-Foundland) and northern parts of the United States from west to east, southwards to about 45° N (states of Washington, Idaho, South Dakota, Minnesota, Wisconsin, Indiana and Mas- sachusetts) (until now referred to as salinarius and subguttatus). Remarks Type-material of the nearctic species subguttatus sensu Roberts and salinarius Wallis has been exam- ined and in my opinion both are conspecific with fulvus. For salinarius that possibility was already noticed by Holmen (1987). Crotch (1873) menti- oned subguttatus referring to a Leconte manus- cript, but he did not regard it as a separate species, but as a form of cribarius Leconte. Blatchley (1910) described it as a subspecies of cribarius. Roberts (1913) discovered that subguttatus was not conspe- cific with cribarius and described it as a separate species. His holotype is probably in AMNH. Two of his paratypes have been examined (CNCI). Material examined. - 712 ex. - Austria: 36 ex. (RMNH, NHMW, ZMAN, OLML). — Belgium: 7 ex. (RMNH, RIN, Jon- gema). — Canada: 3 ex. (holotype and 2 paratypes of salinarius); 16, (paratype of subguttatus), Fraz. Val. B.C.; 1 8, (paratype of subguttatus), Antigonish, N.S. (CNCI); 44 ex. (CNCI, Vondel). - Czechoslovakia: 2 ex. (NHMW, OLML). — Denmark: 1 ex. (NHMW). - Finland: 10 ex. (RMNH, ZMAN, RMNH, NHMW). — France: 30 ex. (RMNH, ZMAN, RMNH, NHMW, BMNH, OLMI, Vondel). - Germany: 39 ex. (RMNH, NHMW, ZMAN, BMNH, OLML). - Great Bri- tain: 94 ex. (CNCI, ZMAN, BMNH, Vondel). - Greece: 1 ex. (OLML). - Hungary: 2 ex. (NHMW, ZMAN).-Ireland: 18 ex. (BMNH). - Italy: 3 ex. (CNCI). — Netherlands: 228 ex. (BMNH, ZMAN, FNML, NHMR, NHMW, RIN, RMNH, Cuppen, Drost, Hielkema, Jongema, Onder de Linden, Vondel). — Norway: 38 ex. (RMNH, ZMAN, BMNH, OLML, MZLU, RMNH); 1 8, (lectotype of fulvus var. sparre-schneidert), 19, (paralectotype of fulvus var. sparre-schneidert), Bjer- keng, ix.1878, Schneider, Haliplus Sparre-Schneideri n. sp., fulv. Schneideri det. Ganglbauer, Typus (NHMW). — Poland: 12 ex. (NHMW). — Rumania: 2 ex. (NHMW). — Spain: 1 ex. (Cuppen). - Sweden: 11 (NHMW, ZMAN, VAN VONDEL: Palaearctic Liaphlus RMNH, BMNH). — USA: 1 ex., Larive Lake, South Dakota, 22.vi.1940 (CNCI). - USSR: 1 ex. (BMNH). — Yugoslavia: 3 ex. (CNCI, OLML). - Locality not traced or unknown: 118 ex. (NHMW, BMNH, MZLU, RMNH, OLML), 1 ex., (9 lecto- type of interpunctatus), (BMNH); 1 ex. (lectotype of lap- ponum) (MZLU). 14. Haliplus gafnyi sp. n. (figs. 132-140) Type material: Holotype 4, Israel, Nesher quarry, 27.1.1989, R. Ortal, IES 5344 (Huy). Paratypes: 1 6, Israel, Berekhat, Nesher quarry, 26.xii.1989, R. Ortal, IES 5502 (Angus); 1 &, Cyprus, Zakaki Marshes, ii.1948, G. Mavromoustaki (Angus); 1 6, Syria, Damascus (MNHN). Diagnosis This species resembles abbreviatus and villiersi in a number of characters, but can be distinguished from them by the flat, strongly and densely punc- tured metasternal process. Description Length 2.9-3.1 mm, width 1.5-1.7 mm. Body oblong oval, widest in the middle (fig. 132). Head. - Yellow-brown, vertex brown, moderate strongly punctured, on the vertex strong darkened punctures. Distance between the eyes 1.4-1.6 X eye-width. Antennae (fig. 133) and palpi yellow to yellow-brown. Pronotum. - Yellow to yellow-brown, anterior margin darkened in the middle. Lateral borders straight, finely bordered. Moderate strongly punc- tured, along the base with a partly double row of strong punctures, discal punctures brown, basal punctures black. Elytra. - Yellow, darkened suture narrow, at most reaching the secondary punctures on the disc, vague brown blotches on the shoulders and in the apical part (fig. 132). Primary punctures strong, 28 punctures in the first row. Secondary punctures strong. All punctures darkened (fig. 134). Ventral side. - Yellow to yellow-brown, elytral epipleura yellow, legs yellow-brown, some darken- ing towards the coxae. Prosternal process flat, moderate strongly punctured, narrowed near the coxae (fig. 136), lateral plicae complete (fig. 137). Metasternal process flat with a small pit in the middle, strongly and densely punctured (fig.136). Setiferous striole % X the tibia length, with about 10 punctures, longer of the two tibial spurs 24 X the length of the first tarsal segment (fig. 135). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 138- 140. 105 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 132-140. Haliplus gafnyi, holotype. - 132, dorsal view; 133, antenna; 134, elytral punctuation; 135, dorsal side of hind tibia; 136, prosternal process; 137, lateral view of prosternal process; 138, left paramere; 139, penis; 140, right paramere. 106 Etymology. - Named after the late Mr. Gabriel Gafny, Hydrologist of the Nature Reserve Author- ity of Israel. Biology In temporary pool. Distribution (fig. 303) Only known from Israel. 15. Haliplus guttatus Aubé (figs. 141-149) Haliplus guttatus Aubé, 1836: 27. Lectotype & (here de- signated), unlabelled (MNHN, coll. SEE Aubé) [examined]. Haliplus pelopis Sahlberg, 1907: 21. Lectotype & (here designated), Elis, J. Sahlb., Spec. typ., 2284, Mus. Zool. H.fors, Spec. typ. No. 852, Haliplus pelopis J. Sb. [aedeagus missing] (UZMH) [examined]. Syn. n. Haliplus guttatus; Aubé 1838: 15, Miiller 1900: 113, Ap- felbeck 1904: 359, Fiori 1904: 195, Escalera 1914: 57, Zimmermann 1920: 308, 1924: 135, Guignot 1928: 145, 1933: 232, 1947: 52, 1959: 36, Scholz 1929: 15, E. Burmeister 1939: 213, E. Burmeister 1987: 177, Csiki 1946: 560, Zaitsev 1953: 72, Lagar Mascaro 1968: 73, Ienistea 1978: 294, Franciscolo 1979: 122. Haliplus pelopis; Zimmermann 1920: 314, 1924: 135, Burmeister 1939: 214, Zaitsev 1953: 72, Ienistea 1978: 294. Diagnosis Very variable in shape, from oval to strongly parallel, maculation not always clearly visible. Description. Length 3.6-4.1 mm, width 1.7-2.1 mm. Body parallel to subparallel (fig. 141). Head. - Yellow-red to rust-coloured, dark blotch on vertex, weakly, behind the eyes more strongly punctured. Distance between the eyes 1.7-1.9 X eye-width. Antennae (fig. 142) and palpi yellow- red. Pronotum. - Yellow to rust-coloured. Lateral borders convex to nearly straight, sometimes slightly concave before the basal corners, finely bordered. Weakly and sparsely, in the basal part densely punctured, basal row of darkened punc- tures twice as wide as the elytral punctures. Elytra. - Yellow with a variable pattern of vague, sometimes hardly visible blotches, in most cases a large blotch on the suture (fig. 141). Fine primary punctures basally stronger, about 37 punctures in the first row. Secondary punctures weak. All punc- tures darkened (fig. 143). Ventral side. - Body yellow to yellow-red, elytral epipleura yellow, legs yellow-red, some darkening towards the coxae. Prosternal process flat, slightly VAN VONDEL: Palaearctic Liaphlus narrowed near the coxae (fig. 145), lateral plicae complete (fig. 146), coarsely and densely punc- tured. Metasternal process flat with a shallow pit in the middle, weakly and sparsely punctured (fig. 145). Setiferous striole on dorsal face of hind tibia about 4 X the tibia length with 5-10 punctures, longer of the two tibial spurs % to % X the length of the first tarsal segment (fig. 144). Length of hind tarsus including claws 1.3-1.4 X the tibia length, first tarsal segment about 1.3 X the length of the second segment. Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 147-149. Biology In stagnant water, fresh or brackish, even in brooks. Distribution (fig. 297) Mediterranean, south and south-west of France, Corsica, Spain including Baleares, Italy including Sardinia and Sicily, Malta, Yugoslavia, Greece, North Africa: Morocco, Algeria, Tunisia. All re- cords from Poland and the southern states of the USSR are suspect and likely concern other species such as fulvus, variegatus or maculatus. Remarks The types of guttatus and pelopis have been examined. They do not show all differences used in earlier identification-keys. I regard both conspe- cific. Material examined. - 67 ex. - Algeria: 5 ex.; 2 ex., Algiers, v.1861, H. Clark; 3 ex., Kraatz; 3 ex., lac des Serpents, Djijellé, Massif des Mouzaia (BMNH); 1 ex. Philippeville (MNHN); 2 9, Taguin; 4 ex., Prov. d’Alger, Taguin, 1895, de Vauloger (NHMW). — France: 1 ex., Nizza, (NHMW); 1 ex., Montpellier, v.1861, H. Clark (BMNH); 1 ex., Camargue, nr. Salin de Giraud, 12.vi.1980 (Huijbregts); 1 ex., Albaron (B. du R.), L. Schaefer (RMNH); 1 ex., Albaron (B. du R.); 1 ex., Pyrenées; 1 ex., Montfort, Landes (ZMAN); 2 ex., Beragne (MNHN); | ex., Les Angles, Gard. iv.1928; 1 ex., Camargue; 1 ex., Corsica, 1905, Aleria; 4 ex., Hyères (BMNH). - Greece: 1 & (Lec- totype of pelopis) (UZMH); 1 ex. (MNHN); 2 ex., Acarnan, 8.1.1969, Forster (OLML); 1 ex., Attika (ZMAN); 4 ex., Zante, Kalamaki, 1909, M. Hilf; 1 ex., Khalkidhiki, Sitho- nia, 22.vii.1988, M Jäch; 1 ex., Kérkira, Stephanides (NHMW). — Italy: 1 ex., Sardinia, Assuni, Krausse (NHMW); 1 À, Sardegna, Gag., 7 km W. of Carbonia, alt. 20 m., 6.v.1984 (Huijbregts); 1 ex., Lucania, Policorof M.T., 3.xi.1972, Angelini; 1 ex., Toscane, Dahl (RMNH); 1 6, Florenz, Brozzi (OLMI); 1 ex., Etruria, (BMNH). — Malta: 1 ex., Chadwick Lakes, 31.1.1987, M. Borg, fresh water (NHMW). - Spain: 2 ex., Palencia (MNHN, ZMAN). — Yugoslavia: 1 &, Zadar (Zara), Sturany, 1891 (NHMW). - Locality not traced: 1 ex., S. Sparato, Dodero (MNHN) 107 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 141-149. Haliplus guttatus, Zante, Greece. - 141, dorsal view; 142, antenna; 143, elytral punctuation; 144, dorsal side of hind tibia; 145, prosternal process; 146, lateral view of prosternal process; 147, left paramere; 148, penis; 149, right paramere. 108 - Without locality: 1 6, Lectotype of guttatus; 1 9, 2 8, Paralectotypes of guttatus (MNHN); 2 ex., 67-56 (BMNH); 1 ex., UlIr.; 2 ex., Dhl., aa 44 (NHMW) 16. Haliplus holmeni sp. n. (figs. 150-160) Type material: Holotype g, [China] “Yunnan, Museum Paris, coll. Maurice Régimbart, 1908’ (MNHN). Diagnosis Looks related to sharpi, but the male has a left paramere with a solid digitus. The fore tarsi have a cushion between the claws. Description Length 3.3 mm, width 1.7 mm. Body oval, widest in the middle (fig. 150). Head. - Brown with yellow labrum, dark blotch on vertex, moderately strongly punctured, unpunc- tured band between the hind parts of the eyes. Distance between the eyes 1.2 X eye-width. Anten- nae yellow, base of eighth till eleventh segment slightly darkened, top of ultimate segment pale (fig. 151), palpi yellow. Pronotum. - Yellow-brown to brown anteriorly, lateral borders somewhat irregularly convex, brown and clearly bordered. Width at the base 2.0 the length in the middle and 1.9 X the width in front. Sparsely, anteriorly densely, basally strongly punctured. Basal punctures black and partly wi- dened. Slightly impressed along the central part of the base (fig. 150). Elytra. - Yellow-brown with extensive black markings: wide band along the base till the fifth puncture-row, suture, arrow-shaped central blotch and blotches on the apical half, the apex and three large blotches on the fifth to ninth interval. Prim- ary puncture-rows strong, about 24 punctures in the first row, third to fifth row close to each other in the central part. Secondary punctures moder- ately strong (fig. 152). Primary and secondary punctures tending to form transverse impressions in the basal part. All punctures darkened. Ventral side. - Rust-coloured to brown, elytral epipleura yellow-brown, legs yellow-red, darkened towards the coxae. Prosternal process slightly im- pressed, stronger impressed apically, densely punc- tured, slightly narrowed near the coxae (fig. 155), lateral plicae complete (fig. 156). Metasternal pro- cess flat, impressed apically and a shallow pit in the middle, sparsely punctured (fig. 155). Coxal plates rather strongly punctured, along the suture weakly punctured, short hairs in the punctures, only few small punctures between the normal ones (fig. 157). Hind margins of the fifth and sixth abdom- inal sternite with complete puncture-rows (fig. VAN VONDEL: Palaearctic Liaphlus 157). Setiferous striole consisting of about 6 separ- ate punctures, longer of the two tibial spurs about % the length of the first tarsal segment (fig. 154). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Between the fore-claws a small cu- shion (fig. 153). Penis and parameres as in figs. 158-160, left paramere with a parallel solid digitus. Etymology. - This species is named after my colleague specialist of Haliplidae, Mogens Hol- men. Distribution (fig. 312) Only known from south-west China (Yunnan) 17. Haliplus jaechi sp. n. (fig. 161-169) Type material: Holotype 6, [Turkey], ‘TR, Bismil 6, Diyarbakir, 29.v.1987 (35), Jäch’ (NHMW). Diagnosis This species is related to kulleri and ortali. The left paramere has a strongly developed solid digitus and on the inner margin a well developed row of setae. Description: Length 3.3 mm, width 1.6 mm. Body oblong oval, widest in the middle (fig. 161). Head. - Brown, puncturation moderately strong and dense, unpunctured band between the hind parts of the eyes. Distance between the eyes 1.4 X eye-width. Antennae yellow-brown, second to fourth segment yellow, third segment 1.5 X the length of the fourth segment (fig. 162). Palpi yel- low-brown. Pronotum. - Yellow-brown. Lateral borders slightly convex, darkened and clearly bordered. Width at the base 1.8 X the length in the middle and 1.5 X the width in front. Sparsely punctured, on the disc almost unpunctured, along the base some slightly darkened larger punctures. Along the base weakly impressed (fig. 161). Elytra. - Yellow to yellow-brown, darkened su- ture and apex, vague blotches on the even intervals. Near the base a transverse slightly darkened band. Primary puncture-rows moderately strong, 28 punctures in the first row. Secondary punctures moderately strong, accompanied by very small punctures (fig. 163), all punctures darkened. Ventral side. - Yellow to yellow-brown, elytral epipleura yellow, legs yellow-brown, darkened to- wards the coxae. Prosternal process slightly con- vex, strongly narrowed near the coxae, rather weakly punctured (fig. 165), lateral plicae complete 109 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 150-160. Haliplus holmeni, holotype. - 150, dorsal view; 151, antenna; 152, elytral punctuation; 153, fore claws; 154, dorsal side of hind tibia; 155, prosternal process; 156, lateral view of prosternal process; 157, hind coxal plate; 158, left paramere; 159, penis; 160, right paramere. 110 VAN VONDEL: Palaearctic Liaphlus £ yy EH) ; SOR DO LOTO. COLI . » "o © © © © 00000 6 © ® © » > a © 290.0 9 © © os OO AUR GU AE. 00 Es Ge Ge © Crag oo 6 oe o 0 TO CR DEE O e + , holotype. - 161, dorsal view; 162, antenna; 163, elytral punctuation; 164, dorsal side , prosternal process; 166, lateral view of prosternal process; 167, left paramere; 168, penis; 169, right Figs. 161-169. Haliplus jaechi of hind tibia; 165 paramere. iat TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 170-178. Haliplus kotoshonis, Okinawa. - 170, dorsal view; 171, antenna; 172, elytral punctuation; 173, dorsal side of hind tibia; 174, prosternal process; 175, lateral view of prosternal process; 176, left paramere; 177, penis; 178, right paramere. 112 (fig. 166). Metasternal process flat with a shallow impression in the middle, sparsely punctured (fig. 165). Coxal plates sparsely punctured. Setiferous striole on dorsal side of hind tibia about ‘4 of the tibia length, with about 7 punctures, longer of the two tibial spurs % the length of the first tarsal segment (fig. 164). Male sexual characters. — First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 167- 169, left paramere with a solid digitus and a well developed row of long bristles on the inner margin. Etymology. — Named after the collector Dr. Manfred Jäch. Distribution (fig. 302) Only known from south-east Turkey, Bismil. 18. Haliplus kotoshonis Kano & Kamiya (figs. 170-178) Haliplus kotoshonis Kano & Kamiya, 1931: 2. Types burned during World War II (Nakane pers. comm.). Haliplus kotoshonis; Sato 1984: 2, Nakane 1985a: 64, 1987: 30. Diagnosis This species resembles pulchellus and maculi- pennis by having the eyes close to each other, but it can be distinguished by the simply impressed prosternal process and the lack of a basal black elytral band. Description Length 3.5-3.6 mm, width 1.9-2.0 mm. Body oval, middle part subparallel, widest in the middle (fig. 170). Head. - Brown, along the eyes dark brown. Puncturation strong and dense. Clypeus impressed in the middle. Distance between the eyes 0.8-1.0 X eye-width. Antennae (fig. 171) and palpi brown- red. Pronotum. - Brown-red. Lateral borders slightly convex, finely bordered. Hind corners protruding and strongly rounded. Punctures very strong, ex- cept on the disc dense and along the base often coarse. Base strongly impressed. Elytra. - Yellow-brown, dark brown maculation on sutural interval and on the apex, a central blotch on the disc and small sometimes connected blotches on the other intervals. Suture slightly bor- dered in the basal part and at the apex. All primary punctures strong to very strong, basal punctures coarse, 32-34 punctures in the first row. Secondary punctures strong and dense. All punctures dar- kened and with a small yellow centre (fig. 172). VAN VONDEL: Palaearctic Liaphlus Ventral side. - Body brown, elytral epipleura yellow-brown, legs yellow-brown to brown, slightly darkened near the coxae. Prosternal pro- cess clearly impressed in the middle, narrowed near the front coxae (fig. 174), lateral plicae complete (fig. 175), strongly and partially coarsely punc- tured. Metasternal process impressed and with a pit in the middle, strongly punctured (fig. 174). Setiferous striole on dorsal face of hind tibia about 4, X the tibia length, containing about 8 punctures, longer of the two tibial spurs about 24 X the length of the first tarsal segment (fig. 173). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 176-178. Distribution (fig. 310) Taiwan (type-locality: isle of Kotosho), Japan (Okinawa). Remarks Although this species seems to belong to the oriental fauna, it is treated here for practical rea- sons. Material examined. - Japan: 1 8, 1 9, Kimibaru, Oh- gimi V., Okinawa-Hontô, 18.viii.1989, leg. E. Matsui (in authors coll.). 19. Haliplus kulleri Vondel (figs. 179-187) Haliplus kulleri Vondel, 1988: 97. Holotype @, Israel, 25.3.81, Lower Galilee, Parod Res., N. Ammud Basin, 35 km W. of Akko, IES 2350, Col. 1673 (NHMW) [examined]. Diagnosis Closely related to jaechi and ortali, but in the males characterised by the long parallel-sided solid digitus on the left paramere. Description Length 2.9-3.0 mm, width 1.6 mm. Body fairly narrow, subparallel, widest in the middle (fig. 179). Head. - Brown, puncturation behind the eyes rather dense, anteriorly more sparse, leaving an unpunctured shiny area between the eyes. Distance between the eyes 1.4-1.5 X eye-width. Antennae (fig. 180) and palpi yellow. Pronotum. - Yellow, slightly darkened on the disc. Lateral borders nearly straight, finely bor- dered. Sparsely and weakly punctured, leaving an unpunctured shiny area on the disc, two short basal rows of slightly widened and darkened punctures (fig. 179). 113 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ® © © OF er . . a 6 e S 0989 9 : nora vedo DO GO OS À > 6 e 6800 8- Figs. 179-187. Haliplus kulleri, holotype. - 179, dorsal view; 180, antenna; 181, elytral punctuation; 182, dorsal side of hind tibia; 183, prosternal process; 184, lateral view of prosternal process; 185, left paramere; 186, penis; 187, right paramere. 114 Elytra. — Yellow with a pattern of brown blotches, one triangular on the middle of the suture and some smaller ones around. Moderately strong darkened primary puncture-rows, about 30 punc- tures in the first row. Secondary punctures small, alternated with very fine punctures (fig. 181). Ventral side. - Body yellow-brown, elytral epi- pleura yellow, legs yellow. Prosternal process slightly convex, narrowed near the coxae, covered with strong punctures (fig. 183), lateral plicae con- tinuous (fig. 184). Metasternal process flat with a shallow pit in the middle, sparsely punctured (fig. 183). Legs half shiny, with fine micro-punctuation. Setiferous striole on dorsal face of hind tibia with about 9 punctures, longer of the two tibial spurs about % X the length of the first tarsal segment (fig. 182). Length of hind tarsus including claws 1.3 X the tibia length, first tarsal segment 1.5 X the length of the second segment. Male sexual characters. - First three tarsal seg- ments of fore- and midlegs hardly widened, scaly hairs on ventral side. Penis and parameres as in figs. 185-187, left paramere with a long parallel digitus (fig. 185). Female: Unknown. Biology In temporary pool. Distribution (fig. 302) Only known from Israel, Lower Galilee. Material examined: 2 ex. - Israel: 1 & (holotype); 1 4, Berekhat Shosh, 23.1.1989, IES 5343 (HUJI). 20. Haliplus laminatus (Schaller) (figs. 188-198) Dytiscus laminatus Schaller, 1783: 314. Types not located, Schaller’s material has been lost (Horn & Kahle 1936) [not examined]. Haliplus cinereus Aubé, 1836: 30. Lectotype (here desig- nated), Gallia, 2645, coll. Chevrolat, Det. Sharp. 82, Haliplus cinereus Aubé p18, affinis Steph., Type, cine- reus Aubé, Type d’Aube, TYPE (pink label) (ISNB) [examined]. Haliplus laminatus ?var. ater Redtenbacher, 1842: 8. Types not located [not examined]. Haliplus laminatus Q var. punctulatus Penecke, 1901: 11. Types not located [not examined]. Haliplus lombardus Fiori, 1904: 198. Types not located [not examined]. Haliplus laminatus; Seidlitz 1887: 31, Apfelbeck 1904: 361, Fiori 1904: 197, Zimmermann 1920: 310, 1924: 131, Guignot 1928: 142, 1933: 229, 1947: 48, Kinel 1929: 218, Scholz 1929: 15, Balfour-Browne 1938: 12, 1940: 154, 1951: 5, 1953: 5, Burmeister 1939: 213, Horion 1941: 365, Csiki 1946: 559, Zaitsev 1953: 67, Gueorguiev 1958: 44, 1960: 23, Lindroth 1960: 42, VAN VONDEL: Palaearctic Liaphlus Lagar Mascaro 1968: 71, Freude 1971: 15, Seeger 1971a: 430, 1971b: 558, 1971c: 166, Ienistea 1974: 207, 1978: 294, Ali 1976: 92, Galewski 1976: 30, Mate- leshko 1977: 68, Franciscolo 1979: 113, Fichtner 1981:327, Holmen 1981: 2, 1987: 110, Leblanc 1984: 8, Lundberg 1986: 11, Vondel 1986: 128, 1989a: 57. Haliplus cinereus, Aubé 1836: 30, 1838: 18, Erichson 1837: 185, Zimmermann 1920: 310, Balfour-Browne 1936: 75, 1938: 12, Burmeister 1939: 213. Haliplus laminatus ?var. ater; Zimmermann 1920: 310. Haliplus Q var. punctulatus; Zimmermann 1920: 310. Haliplus lombardus, Zimmermann 1920: 312, Burmeis- ter 1939: 213. Diagnosis This species can be distinguished from all other palaearctic species of Liaphlus by the dark lines on the elytral puncture-rows and in the males by the strongly dilated first mesotarsal segment and the lack of a digitus on the right paramere. Description Length 2.5-3.6 mm, width 1.5-1.9 mm. Body with subparallel sides in the central part (fig. 188). Head. - Yellow-brown with brown collar, with dense and on the collar strong and darkened punc- tures. Distance between the eyes 2.0-2.1 X eye- width. Antennae (fig. 189) and palpi yellow. Pronotum. - Yellow. Basally not as wide as base of elytra. Lateral borders slightly convex, finely bordered. Densely, on the disc more sparsely punc- tured. Elytra. - Yellow to yellow-red, strong primary puncture-rows with widened and usually groove- like basal punctures, about 42-46 punctures in the first row. Secondary punctures rather weak. All punctures darkened (fig. 190). Primary rows and suture partly, sometimes completely covered with dark lines, not connected to each other (fig. 188). Ventral side. - Yellow to yellow-brown, elytral epipleura yellow, legs yellow to yellow-brown, coxae red-brown. Prosternal process grooved, nar- rowed near the coxae, widened towards the apex, punctures moderately strong and dense (fig. 193), lateral plicae complete (fig. 194). Metasternal pro- cess slightly impressed in the middle, moderately strongly punctured (fig. 193). Setiferous striole on dorsal side of hind tibia stretched over the apical %, longer of the two tibial spurs % X the length of the first tarsal segment (fig. 195). Male sexual characters. - First three protarsal segments slightly widened, scaly hairs on ventral side, first segment ventrally with a complete row of strong spines (fig. 191). First mesotarsal seg- ment strongly delated ventrally (fig. 192). Penis and parameres as in figs. 196-198, right paramere without a digitus as characteristic for Liaphlus (fig. 198). Female. - Elytra with strong micropuncturation. 115 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 N O = 191 © SCE © | 5 © so ees y het ete ni 2 » © © © © Sarre. te NAR © oes 29 Bee oe È e ° ; DA O Peony È OOO Foo ia» 0500 er. a ° ° e 090° 00 sastre ° ° 5 2 > OOAD 00000 e 00080 ° nrd a 0 000290» ere. rf 0 . = © © © 0 mmm © ER = ar 7 grace © © © e one? 3 DG eo + se at 4, lateral view of prosternal ; 189, antenna; 190, elytral punctuation; 191, first ; 197, penis; 198, right paramere. ; 193, prosternal process; 19 B 2) > = 58 kl ot Al ap OEH oo. U —oE I ns ves e va Ss Cw da u ao WA os aM «a nn DER OP a og .« “75 Qu en = B= RE, (N et Figs. 188-198. Haliplus laminatus process; 195, dorsal side of hind tibia tarsal segments of forelegs 116 Biology Usually in slow running water of rivers, canals and brooks, but also found in silt ponds. The adults feed on filamentous algae and invertebrates like larvae of Chironomidae and Oligochaeta (Seeger 1971b) and freshly killed Asellus aquaticus (Vondel 1986). The larvae (3rd and 2nd instar) were des- cribed by Vondel (1986). Distribution (fig. 293) Central Europe from Denmark to the east coast of Spain, central Italy and Albania and from south- east England to the European part of the USSR. The species is also recorded from Iraq (Ali 1976). The records from Ireland (Burmeister 1939, Fran- ciscolo 1979) are doubtful. Remarks This species is usually treated as belonging to the subgenus Liaphlus, although Kinel (1929) erected a separate subgenus Hoplites for this species. The name Hoplites is preoccupied (see: International Commission on Zoological Nomenclature 1955: opinion 353), but never replaced. There is reason to believe that this species is more closely related to the subgenus Haliplus s. str. (Vondel 1986; Hol- men 1987). Before erecting new subgeneric names it seems better to me to await the results of syste- matic studies on a world basis. Material examined. - 481 ex. - Albania: 2 ex. (NHMW). — Austria: 81 ex. (ZMAN, CNCI, NHMW, NHMR). - Belgium: 2 ex. (RMNH). — Czechoslovakia: 18 ex. (ZMAN, CNCI, RMNH, MHNG, NHMW, RMNH). - France: 12 ex. (NHMW, RMNH, Jongema, BMNH, Vondel). — Germany: 20 ex. (ZMAN, NHMW, RMNH). — Great Britain: 5 ex. (BMNH). — Hungary: 3 ex. (NHMW). - Italy: 12 ex. (RMNH, NHMW). — Luxembourg: 1 ex. (ZMAN). - Netherlands: 291 ex. (ZMAN, RIN, RMNH, Cuppen, Drost, Giessen, Huijbregts, Jongema, Vallenduuk, Vondel). - Poland: 11 ex. (ZMAN, RMNH, NHMW). — Spain: 2 ex. (NHMW). - Locality not traced or unknown: 22 ex. (BMNH, NHMW, RMNH); 1 ex. (lectotype of cinereus) (ISNB). 21. Haliplus maculatus Motschulsky (figs. 199-207) Haliplus maculatus Motschulsky, 1860: 100. Lectotype 9 (here designated), [USSR, n.w. of Caspian sea], ‘Bet- scho-Ko’ [red label, hardly readable] ‘Haliplus macu- latus Motsch., Des K. occ., 1295’ [red label] (ZMUM) [examined ]. Haliplus syriacus Wehncke, 1880: 73. Lectotype 9 (here designated) ‘Syria, Sharp’ (MNHN) [examined]. Syn. n. Haliplus zimmermanni Gschwendtner, 1921: 56. Lecto- type & (here designated) ‘Oberösterreich, L. Gschwendtner, Umg. Linz, 5.5.1909, Type, Coll. Gschwendtner, Haliplus Zimmermanni Gs, det. VAN VONDEL: Palaearctic Liaphlus Gschwendt, Type’ [examined]. Syn. n. Haliplus maculatus; Zimmermann 1920: 317, Kolossov 1931: 116, Zaitsev 1953: 75, Ienistea 1978: 294. Haliplus syriacus; Zimmermann 1920: 317, 1924: 136, Guignot 1933: 244, Zaitsev 1953: 74, Burmeister 1985: 41. Haliplus zimmermanni; Zimmermann 1924: 134, Müller 1926: 167, Guignot 1933: 241, Horion 1941: 365, lenistea 1978: 295, Franciscolo 1979: 120. [aedeagus missing] (OLML) Diagnosis This species closely resembles small specimens of fulvus. The male can be distinguished from fu/- vus by the narrow and somewhat pointed top of the penis. Description Length 3.4-4.0 mm, width 1.8-2.2 mm. Body oval, widest in the middle (fig. 199). Head. - Yellow-brown to rust-coloured, vertex usually slightly darkened, weakly to moderately punctured, unpunctured band between the eyes. Distance between the eyes 1.4-1.6 X eye-width. Antennae yellow, first segment brown (fig. 200), palpi yellow. Pronotum. - Yellow to rust-coloured. Lateral borders straight, finely bordered. Sparsely and weakly, along the base strongly punctured, basal punctures usually slightly darkened (fig. 199). Elytra. - Yellow to rust-coloured, rather weak to moderately strong primary puncture-rows, 28-38 punctures in the first row. Secondary punctures moderately strong, accompanied by very small punctures (fig. 201), usually all punctures dar- kened. Markings variable, oblong blotches along the suture, on the apex and on the even intervals, the central blotches on the second and fourth inter- val often connected anteriorly, the markings are often hardly visible or even absent (fig. 199). Ventral side. - Yellow-brown to rust-coloured, elytral epipleura yellow to yellow-red, legs yellow- red to rust-coloured, darkened towards the coxae. Prosternal process slightly convex, rather sparsely punctured, narrowed near the coxae (fig. 203), lat- eral plicae complete (fig. 204). Metasternal process flat with a shallow pit in the middle, sparsely punc- tured (fig. 203). Coxal plates with moderately strong and in between fine punctures, the strong punctures absent on a wide band along the suture. Setiferous striole on dorsal side of hind tibia about % of the tibia length, with 7-10 punctures, longer of the two tibial spurs about % the length of the first tarsal segment (fig. 202). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 205- 207. 117 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 e e Figs. 199-207. Haliplus maculatus, Vienna, Austria. - 199, dorsal view; 200, antenna; 201, elytral punctuation; 202, dorsal side of hind tibia; 203, prosternal process; 204, lateral view of prosternal process; 205, left paramere; 206, penis; 207, right paramere. 118 Biology In temporary pools, in brooks. Distribution (fig. 300) Austria, Hungary, Czechoslovakia, Rumania, Yugoslavia, Italy? (identification uncertain), Po- land?, Israel, Syria, Afghanistan, USSR near border with Iran east of the Caspian sea and north of the Black and the Caspian sea. All European specimens I examined are from more than 60 years ago, so the species might be extinct in Europe. Remarks The types of maculatus, syriacus and zimmer- manni have been compared. Although the first two are females and the important character of the pointed top of the penis cannot be checked, I con- sider the three conspecific. Material examined. — 41 ex. - Afghanistan: 1 9, W. Afghanistan, Hari Rud, Herat, 20.vii.1949, Danish Cen- tral Asian Expedn, St. no. 4, Ex stomach of Lobivanellus indicus, Haliplus fulvus, J. Balfour-Browne det. 1964 (BMNH). — Austria: 1 & (lectotype zimmermanni); 1 9, Wien, Prater, Kniz (OLML); 2 6, Neusiedler See; 1 4, Wien surr., Smolik; 2 9, Lavant Tal, Car (RMNH); 1 @, Gars, Minarz; 5 6, 1 9, Wien, Prater; 1 @, surr. Wien; 1 6, 2 9, Steiermarken (Styria), Kars; 1 6, Klosterneuburg, Pinker (NHMW). — Czechoslovakia: 1 &, Veseli (Moravia, surr. Ostra), Th. v. Wanka (NHMW). - Hungary: 2 6, Ujpest (RMNH); 1 @, Szekesfehervar (Szfehervar), 1928, (OLML). — Israel: 1 6, Berekhat Sasa, 10.viii.1985, Ortal & Jäch (Hujr); 2 9, Jerusalem, Sauliy (MNHN). — Italy: 1 ©, Toscana, Dahl (RMNH). - Rumania: 1 4, Bocsa Vasovei (N-Bogsan, Banat) (RMNH). - Syria: 1 9 (lectotype) (MNHN). — Yugoslavia: 1 9, Fruska Gora (NHMW). - USSR: 1 Q (lectotype of maculatus) (ZMUM); 1 9, Kopet- Dagh, Germab (OLML); 2 9, Geok Tepe (Caucase, Guéox- Tapa), L. Mesmin (Angus); 1 6, Geok Tepe (Caucase, Gok Tepe), L.Mesmin (OLMI). - Locality not traced: 1 4, Munganast, Type,1921, coll. Gschwendtner, Haliplus Zimmermann Gschw., det. L. Gschw., Paralectotype [aedeagus missing] (OLML). - Without locality: 2 3, (OLML); 1 @, Ullr. (NHMW). 22. Haliplus maculipennis Schaum (figs. 208-216) Haliplus maculipennis Schaum, 1864: 107. Haliplus maculipennis; Régimbart 1877: 354, Sahlberg 1913: 40, Zimmermann 1920: 312, 1924: 138, Guignot 1959: 37, Alfieri 1976: 38. Diagnosis This species can be distinguished from other palaearctic species by the distance between the eyes being about equal to eye-width and the row of dark blotches on the elytral marginal parts. VAN VONDEL: Palaearctic Liaphlus Description Length 3.0-3.3 mm, width 1.6-1.7 mm. Body wide with subparallel sides (fig. 208). Head. - Yellow-red with darker vertex, weakly to moderately strongly punctured. Distance between the eyes about eye-width. Antennae (fig. 209) and palpi yellow-red. Pronotum. - Yellow-red. Lateral borders straight, strongly bordered, posterior corners usu- ally protruding. Strong punctures along base (fig. 208). Elytra. — Yellow-red, primary puncture-rows weak except in the basal part of the first 3 or 4 rows, basal part of third row usually impressed, about 30 punctures in the first row. Secondary punctures weak (fig. 210). Extensive elytral mark- ings: complete suture, a basal band until the 5th puncture-row, triangular apically hooked central blotch and several smaller blotches, 8th puncture- row with a row of small blotches (fig. 208). Ventral side. - Body yellow-red, legs yellow-red, darkened towards the coxae. Prosternal process wide, slightly narrowed near the coxae, strongly impressed towards the apex, sparsely but strongly punctured (fig. 212), lateral plicae complete (fig. 213). Metasternal process flat with a pit in the middle, weakly punctured (fig. 212). Setiferous striole about Y, X the tibia length, longer of the two tibial spurs about *4 X the length of the first tarsal segment (fig. 211). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 214- 216. Distribution (fig. 306) Egypt. The variety abyssinicus Régimbart in Egypt, Sudan, Ethiopia, Zaire, Ivory Coast and Bénin. Subspecies capensis Gueorguiev in South Africa. Remarks In the collection of München (ZSMC) 1 Q is pres- ent indicated as type: Egypt, Schaum, Typ, Samm- lung Cl. Miiller, Haliplus maculipennis Schaum [examined]. According to Guignot (1959) the type should be in ZMHB. [not examined]. Also because this species originates from the Ethiopian fauna I postponed the designation of a lectotype and the examination of the described varieties or subspe- cies until a revision of the Ethiopian fauna is un- dertaken. Material examined. - 4 ex. - Egypt: 1 9, Schaum, Typ (ZSMC); 1 g, Cairo, J.Sahlb. (NHMW).- Without locality: 1 9, 18 (ZSMC). 119 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 208-216. Haliplus maculipennis (208, 212-216, locality unknown; 209 -211, Cairo, Egypt). - 208, dorsal view; 209, antenna; 210, elytral punctuation; 211, dorsal side of hind tibia; 212, prosternal process; 213, lateral view of prosternal process; 214, left paramere; 215, penis; 216, right paramere. 120 23. Haliplus mucronatus Stephens (figs. 217-225) Haliplus mucronatus Stephens, 1829: 40. Types probably in BMNH [not examined]. Haliplus badius Aubé, 1836: 25. Types probably in MNHN [not examined]. Haliplus parallelus Babington, 1836: 178. Types possibly in CUMZ [not examined]. Haliplus siculus Wehncke, 1883: 145. Types probably in MNHN [not examined]. Haliplus mucronatus, Schaum 1848: 42, Miiller 1900: 113, Apfelbeck 1904: 359, Fiori 1904: 193, Escalera 1914: 57, Zimmermann 1920: 312, 1924: 131, Guignot 1928: 143, 1933: 229, 1947: 49, 1959: 33, Scholz 1929: 14, F. Burmeister 1939: 213, Balfour-Browne 1940: 149, 1953: 5, Horion 1941: 365, Bertrand 1949: 25, Zaitsev 1953: 67, Gueorguiev 1958: 44, 1981: 400, Lagar Mascaro 1968: 72, Freude 1971: 15, Franciscolo 1972: 65, 1979: 115, Ali 1976: 92, Ienistea 1978: 294, Fichtner 1981: 327, Leblanc 1984: 40, Vondel 1985: 33, E. Burmeister 1987: 177, Holmen 1987: 115. Haliplus badius; Aubé 1838: 13, Schaum 1848: 42, Séidlitz 1887: 29, Zimmermann 1920: 312. Haliplus parallelus, Schaum 1848: 42, Zimmermann 1920: 312. Haliplus siculus; Seidlitz 1887: 29, Apfelbeck 1904: 359, Scholz 1916: 165, Zimmermann 1920: 312. Diagnosis This species can be distinguished from related species by the large distance between the eyes (more than 2.3 X eye-width) and the metasternal - process, not being grooved or pitted in the middle. Description Length 4.0-4.4 mm, width 1.9-2.2 mm. Body subparallel (fig. 217). Head. - Yellow-red to brown, punctures moder- ately strong. Distance between the eyes 2.3-2.8 X eye-width. Antennae (fig. 218) and palpi yellow- red. Pronotum. - Yellow-red. Lateral borders slightly convex, weakly tapering anteriorly, finely bor- dered. Densely and fairly strongly punctured, ba- sally with a row of widened punctures, on the disc with an unpunctured transverse band. Punctures usually darkened (fig. 217). Elytra. - Yellow-red without markings, all prim- ary puncture-rows about equal in strength, 45-50 punctures in the first row. Secondary puncture- rows well developed, number of punctures about the same as in the primary rows. All punctures darkened (fig. 219). Basal punctures of 3rd to 5th row usually groove-like. Ventral side. - Body yellow to yellow-red, legs yellow to yellow-red. Prosternal process flat to slightly impressed, strongly, apically even coarsely punctured, gradually narrowed towards the coxae (fig. 221), lateral plicae complete (fig.222). Metas- ternal process slightly convex, sparsely punctured VAN VONDEL: Palaearctic Liaphlus (fig. 221). Setiferous striole on dorsal side of hind tibia long, more than half of the tibia length, longer of the two tibial spurs about two thirds the length of the first tarsal segment (fig. 220). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 223- 225. Biology In fresh or brackish water, particularly in clay and gravel pits with Characeans. The larva was described by Bertrand (1928, as guttatus). Distribution (fig. 294) Central and southern Europe: south and sou- theast of England, the Netherlands, Belgium, France including Corsica, Spain including Mallorca, the south of Germany, Switzerland, Italy including Sardinia and Sicily, Yugoslavia, Albania, Greece, Turkey, USSR along the Black Sea, Iraq, North Africa along the coast from Morocco till Libya. The record from the Harz mountains in East Germany was confirmed by Fichtner (1981), but he is not sure that the labelling is correct. The records from Leningrad and the Kirghiz steppes are incorrect (Zaitsev 1953). Material examined. - 116 ex. - Albania: 4 ex. (NHMW). - Algeria: 1 ex. (BMNH). - France: 26 ex. (NHMW, RMNH, Jongema, ZMAN, Vondel). - Greece: 8 ex. (NHMW, Kanaar, ZMAN). — Italy: 23 ex. (NHMW, CNCI, Jongema, ZMAN, RMNH). - Netherlands: 21 ex. (ZMAN, RMNH, Vondel). — Spain: 22 ex. (NHMW, CNCI, RMNH, ZMAN, Cuppen). — Yugoslavia: 10 ex. (NHMW, RMNH, ZMAN). — Locality not traced or unknown: 5 ex. (RMNH, NHMW). 24. Haliplus ortali sp. n. (fig. 226-234) Type material: Holotype 8, [Turkey] ‘TR, Bismil 6, Diyarbakir, 29.5.1987 (35), Jach’ (NHMw). Paratypes (15 ex.): Israel: 1 @, 15.2.1982, Nesher quarry, rainpool, Haifa, 15.ii.1982, R. Ortal & A. Sofer, IES 2575, Col. 1751- 01; 1 4, Nesher quarry, rainpool near Haifa, 27.1.1989, R. Ortal, IES 5344; 1 8, Golan, Birket sinndiane, 11.ii.1981, R. Ortal, IES 2502, Col. 2744; 1 9, Hula, Shosh pool, 12.1.1982, Z. Kuller & G. Gafny, IES 2599, Col. 1758; 1 6, 2 9, Berekhat, Nesher quarry, 26.xii.1989, R. Ortal, IES 5502; 1 g, Ein Abu Mashag, 12.ix.1989, IES 5401 (HUJI). - Cyprus: 2 6, 4 9, Zakaki Marshes, ii.1948, G. Mavro- moustakis (Angus). — Syria: 1 8, Damascus, brooklet (MNHN). Diagnosis This species resembles very much species like kulleri, jaechi or even abbreviatus, gafnyi and vil- lierst. The males can be distinguished by the 121 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 J Netherlands. - 217, dorsal view; 218, antenna; 219, elytral punc- Figs. 217-225. Haliplus mucronatus, Oostvoorne, : 222, lateral view of prosternal process; 223, left tuation; 220, dorsal side of hind tibia; 221, prosternal process paramere; 224, penis; 225, right paramere. 122 VAN VONDEL: Palaearctic Liaphlus . se eeoses 0 0 ec oe oe? .a Figs. 226-234. Haliplus ortali, holotype. - 226, dorsal view; 227, antenna; 228, elytral punctuation; 229, dorsal side of hind tibia; 230, prosternal process; 231, lateral view of prosternal process; 232, left paramere; 233, penis; 234, right paramere. 123 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ‘Se e e oe o © lio a 0 200 S- (8 => A Figs. 235-243. Haliplus ovalis, holotype. - 235, dorsal view; 236, antenna; 237, elytral punctuation; 238, dorsal side of hind tibia; 239, prosternal process; 240, lateral view of prosternal process; 241, left paramere; 242, penis; 243, right paramere. 124 pointed digitus on the left paramere. In the Turkish holotype that digitus is at the top provided with 3 setae, while the Israeli paratypes have only 1 seta. Description Length 2.9-3.4 mm, width 1.5-1.8 mm. Body oval, widest in the middle (fig. 226). Head. - Yellow-brown to brown, slightly dar- kened on the vertex, weakly punctured, unpunc- tured band between the hind parts of the eyes. Distance between the eyes 1.4-2.0 X eye-width. Antennae yellow (fig. 227), palpi yellow. Pronotum. - Yellow to yellow-brown on the disc, lateral borders straight, darkened and finely bor- dered. Width at the base 2.0 X the length in the middle and 1.6 X the width in front. Weakly and sparsely punctured, on the disc almost unpunc- tured, along the base some larger darkened punc- tures, along the base weakly impressed (fig. 226). Elytra. - Yellow to yellow-brown, apical % of suture darkened, clear discal blotch connected to the dark suture or vague darkening and darkened puncture-rows on the disc and vague blotches in the apical half in the even intervals. Primary punc- ture-rows fairly strong, about 32 punctures in the first row. Secondary punctures moderately strong, accompanied by very small punctures (fig. 228), all punctures strongly darkened. Ventral side. - Yellow to yellow-brown, pro- and metasternal process brown, elytral epipleura yel- low, legs yellow-brown, darkened towards the coxae. Prosternal process slightly convex, nar- rowed near the coxae, moderately strongly punc- tured (fig. 230), lateral plicae sometimes inter- rupted along the front coxae (fig. 231). Metasternal process flat with a shallow impression in the mid- dle, sparsely punctured (fig. 230). Coxal plates rather densely punctured, small punctures between the larger ones, sutural area only weakly and spar- sely punctured. Hind margins of the fifth and sixth abdominal sternite with only few punctures, coxal plates reaching the fifth sternite. Setiferous striole about ‘4 of the tibia length, with about 6 coarse punctures, longer of the two tibial spurs % the length of the first tarsal segment (fig. 229). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side, penis and parameres as in figs. 232-234, left paramere with a pointed solid digitus, at the top provided with 1 to 3 setae. Etymology. - Named after Dr. R. Ortal who collected part of the type-material. Biology In temporary pools and in brooklet. VAN VONDEL: Palaearctic Liaphlus Distribution (fig. 303) South-east Turkey, Cyprus, Syria and Israel, 25. Haliplus ovalis Sharp (figs. 235-243) Haliplus ovalis Sharp, 1884: 440. Lectotype & (here de- signated), ‘Japan, Jokohama, 29.10.81, Lewis, Type, Haliplus ovalis Type D. S., Sharp Coll., 1905-313’ (BMNH) [examined]. Haliplus ovalis; Zimmermann 1920: 314, 1924: 134, Ta- kizawa 1931: 141, Wu 1932: 343, J. Balfour-Browne 1946: 436, Sato 1960: 252, 1984: 3, Ohkura 1962: pl. 64, Nakane 1963: 55, 1985a: 63, 1985b: 40, 1987: 30, Cho 1969: 166, Yoon 1988: 622. Diagnosis This species differs from the closely related chz- nensis in having the distance between the eyes more than 1.4 X the eye-width and in the male by the abruptly bent top of the penis. Description Length 4.1-4.3 mm, width 2.1-2.2 mm. Body oval, widest in the middle (fig. 235). Head. - Yellow-brown, dark spot on vertex, weakly punctured. Distance between the eyes 1.5 X eye-width. Antennae (fig. 236) and palpi yellow- red. Pronotum. - Yellow, lateral borders straight, finely bordered. Densely, on the disc very sparsely punctured, basal punctures widened and darkened, sometimes only slightly. Elytra. - Yellow, moderately strong primary punctures, about 40 in the first row. Secondary punctures numerous. All punctures darkened (fig. 237). Suture, apical point and 9 or 10 spots dar- kened, central spots on first and third interstriae sometimes connected with suture (fig. 235). Ventral side. - Body yellow to yellow-red, elytral epipleura yellow, legs yellow-red, gradually dar- kened towards the coxae. Prosternal process flat, strongly and densely punctured, narrowed near the coxae (fig. 239), lateral plicae complete (fig. 240). Metasternal process flat with a pit in the middle, rather weakly punctured (fig. 239). Setiferous stri- ole on dorsal side of hind tibia almost half of the tibia length, with about 14 punctures, longer of the two tibial spurs two thirds the length of the first tarsal segment (fig. 238). Male sexual characters. - First three tarsal seg- ment of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 241- 243. Distribution (fig. 309) Japan: Kyushu, Honshu. The record of Korea (Takizawa 1931) probably concerns chinensis. The 125 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 244-252. Haliplus pulchellus, lectotype. - 244, dorsal view; 245, antenna; 246, elytral punctuation; 247, dorsal side of hind tibia; 248, prosternal process; 249, lateral view of prosternal process; 250, left paramere; 251, penis; 252, right paramere. 126 records of China (Zaitsev 1953) likely concern cht- nensis. Remarks Types of the very closely related ovalis and chi- nensis have been examined and compared. Unfor- tunately the holotype of chinensis is a female. The shape of the penis of the lectotype and other exam- ined Japanese specimens of ovalis is different from that of the continental specimens I have seen. So far I consider the Japanese specimens to belong to ovalis and the continental specimens to chinensis. J. Balfour-Browne (see Sato 1984) already noticed a difference in continental material and obviously planned to describe Haliplus ovalis ssp. sinicus from Djalantun, but that name is not known to be published and is thus unavailable. Material examined. — 11 ex. — Japan: 1 @ (lectotype) (BMNH); 1 9, Tokio, Ahlwarth (ZMHB); 1 9, surr. Tokio, Bodemeyer (RMNH); 1 @, Murakami, Niigata Pref., 6- 1l.viii.1961, M. Sato; 1 9, Kurokawa, N-Echigo, S.viii.1961, M. Sato (ZMUC); 1 ex, Settsu, 5.v.1931; 1 ex., surr. Tokio, E. v. Bodemeyer; 1 ex., Lewis (SMFD); 1 ex., Yokohama, 28.ix-3.xi.1881, G. Lewis (MNHN); 2 ex., Ichi- nokawa, Aso T., Kumamoto Pref., 28.viii.1986, E. Matsui (Vondel). 26. Haliplus pulchellus Clark (figs. 244-252) Haliplus pulchellus Clark, 1863 : 418. Haliplus oceanicus Régimbart, 1886 : 139. Haliplus pulchellus var. indicus Régimbart, 1899 : 189. Diagnosis This species differs from other species in the area by the small distance between the eyes and the double furrowed prosternal process. Description Length 2.5-3.4 mm, width 1.5-2.0 mm. Body oval, widest just before the middle (fig. 244). Head. - Brown to dark brown, darkened along the eyes and near the antennae, sometimes with a weak arrow-like darkening in the middle. Sparsely and weakly, behind the eyes more strongly punc- tured. Distance between the eyes 0.9-1.1 X eye- width. Antennae yellow to brown-yellow (fig. 245), palpi yellow-red to brown. Pronotum. - Yellow to yellow-red, large median dark blotch that can be reduced to a round blotch on the anterior half with a partly reduced or vague or completely absent blotch on the posterior half (fig. 244). Lateral borders straight to slightly con- vex, anterior corners concave, clearly, vaguely or not visibly bordered, margin never reaching the VAN VONDEL: Palaearctic Liaphlus anterior corner. Apical corners rounded, base near the scutellum usually finely impressed. Strongly, anteriorly weaker punctured, near the basal corners with one or two stronger punctures in a slight depression. Basal punctures darkened. Elytra. - Yellow to yellow-red, clear and ex- tended markings along the suture and along the base until the fifth primary puncture-row, a central blotch in the third interval connected to the suture and various small blotches in the lateral part (fig. 244). Primary punctures of average strength, stronger towards the base, about 30 punctures in the first row. Secondary punctures relatively strong accompanied by some very small punctures. All punctures darkened except the central hole (fig. 246). Lateral borders sometimes slightly serrulate in the basal part. Ventral side. - Body yellow-brown to dark brown, elytral epipleura yellow to yellow-brown, legs yellow-brown with brown to dark-brown femora and coxae. Prosternal process with two strong grooves, bordered anteriorly, punctured, sometimes strongly, especially in the grooves, weakly narrowed near the coxae, slightly diverging towards the apex (fig. 248), lateral plicae complete (fig. 249). Metasternal process flat with two pits, these pits sometimes very strong and than almost forming one confluent pit, sparsely punctured (fig. 248). Coxal plates rather sparsely, near the suture weakly punctured, sutural corners more or less rec- tangular. Sternites with complete puncture-rows. Last sternites almost completely covered with strong punctures, apical point with a very short ridge. Setiferous striole on dorsal side of hind tibia very short, very close to the lateral row of setae and often hardly to distinguish from it, longer of the tibial spurs about % X the length of the first tarsal segment (fig. 247). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 250-252. Distribution (fig. 307) China along the coast north to Xiamen; Thai- land; Vietnam; Indonesia: Sumatra, Java, Sulawesi; India; Sri Lanka. Remarks This species from oriental origin is included in this revision because it penetrates into China, es- pecially along the coast. The species will be treated completely, however, in a revision of the Oriental Haliplidae. Material examined from the treated area: 2 ex., China, 127 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 2 * - 2 ‘ 254 ° OD ii © @ 5 © 5 e ° o Dome Ve spoe eer VERRI 000000 e 090020. cede ee G0 © 90 09° © ©, ‘e @ © E 0 0099 ts 9.0.0. est er: © © LT ‘909000 esse ® Figs. 253-261. Haliplus rubidus, France medit. - 253, dorsal view; 254, antenna; 255, elytral punctuation; 256, dorsal side of hind tibia; 257, prosternal process; 258, lateral view of prosternal process; 259, left paramere; 260, penis; 261, right paramere. 128 Amoy [Xiamen], Galatea (zMUC). The other material will be treated in the revision of oriental Haliplidae. 27. Haliplus rubidus Perris (figs. 253-261) Haliplus rubidus Perris, 1857: 117. Types probably in Ecol. Agricult. Montpellier, France [not examined]. Haliplus perforatus Schaum, 1859: 48. Depository of types unknown [not examined]. Haliplus rubidus, Marseul 1882: 107, Seidlitz 1887: 31, Zimmermann 1920: 314, 1924: 132, Guignot 1928: 143, 1933: 230, 1947: 50, 1959: 35, Scholz 1929: 15, Burmeister 1939: 214, Lagar Mascaro 1968: 72, Ienis- tea 1978: 294, Franciscolo 1979: 119. Haliplus perforatus, Zimmermann 1920: 314. Diagnosis Closely related to flavicollis, from which it differs in the larger punctures on the elytral base, the more pronounced shoulders, the wider distance between the eyes and the usually larger size. Description Length 3.9-4.4 mm, width 2.0-2.3 mm. Body with strong shoulders, tapering strongly behind the middle (fig. 253). Head. - Yellow-red to rust-coloured, weakly punctured. Distance between the eyes 1.7-1.9 X eye-width. Antennae (fig. 254) and palpi yellow- red to rust-coloured. Pronotum. - Yellow-red to rust-coloured. Lateral borders straight, clear darkened margins, narrowed anteriorly. Densely and weakly, basally strongly punctured, some basal punctures widened (fig. 253). Elytra. - Yellow-red to rust-coloured, very strong primary puncture-rows, near the base punctures wider then the distance from each other (fig. 255), about 30 punctures in the first row. Secondary punctures moderately strong, 20 in the second in- terval. Punctures weakly darkened. No markings present. Ventral side. - Yellow-red to rust-coloured, ely- tral epipleura yellow-red, legs yellow-red to rust- coloured, darkened towards the coxae. Prosternal process flat or slightly impressed and very wide apically, coarsely punctured, narrowed strongly near the coxae (fig. 257), lateral plicae absent in front of the coxae (fig. 258). Metasternal process flat, strong pit in the middle, coarsely punctured (fig. 257). Setiferous striole on dorsal side of hind tibia more than half of the tibia length, longer of the two tibial spurs little more than half the length of the first tarsal segment (fig. 256). Male sexual characters. — First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 259- 261. VAN VONDEL: Palaearctic Liaphlus Biology Stagnant water in pools, dune-pools. Distribution (fig. 299) South of France along Atlantic ocean and along the Mediterranean Sea, Italy in the province Ligu- ria, Spain, Portugal, Morocco, Canary Islands. Material examined. - 31 ex. - France: 1 ex., Lac d’Hour- tin, Medoc, 28.ix.1973 (Jongema); 1 ex., Biarritz; 1 ex., Pyrenaei (ZMAN); 2 ex., Gallia m.; 2 ex., Gironde, Vaul- oger; 1 ex., France Merid, Gironde, de Vauloger; 2 ex., F. de la Teste de Vauloger (NHMW); 3 6, 2 9, Pyrenees, Biarritz; 1 @, 2 9, Sauveterre, Gard, v.1928, Guignot; 1 ex.; 1 ex., Landes; 2 ex., Pyrenees (BMNH). - Portugal: 2 ex., Alentejo prov. Rio Mira nr. S. Ana da Serra, 21.ix.1976, no. 60, 37°38'N 8°13'W, v. Berge Hene- gouwen (RMNH); 1 ex., Algarve, Silves, Ribeira de Arade, 10.11.1982; 1 ex, Algarve, Porto de Lagos, barrage, 12.11.1982 (Cuppen). - Morocco: 1 9, Tanger, Haliplus fulvus (MHNG); 1 &, Tanger, vii.1895 (MNHN). — Locality unknown: 1 ex. (BMNH). 28. Haliplus sbarpi Wehncke (figs. 262-270) Haliplus sharpi Wehncke, 1880: 74. Lectotype 9 (here designated), ‘China, Sharp, Dr. Régimbart vidit 1898’ (MNHN) [examined] Haliplus tsukushiensis Yoshimura, 1932: 102. Depository of types unknown [not examined]. Haliplus sharpi; Sharp 1884: 440, Zimmermann 1920: 316, 1924: 137, Takizawa 1931: 142, Wu 1932: 343, Zaitsev 1953: 76, Nakane 1963: 55, 1985a: 63, 1985b: 40, 1987: 30, Sato 1984: 3. Haliplus tsukushiensis, Ohkura 1962: pl. 64, Nakane 1963: 55, 1985b: 40, Sato 1984: 3. Diagnosis This species could be confused with specimens of diruptus in which a basal dark band is present on the elytra. The prosternal process in sharp: is wide and hardly narrowed near the coxae, while diruptus has a narrow prosternal process, which is strongly narrowed near the coxae. In the male the penis of both species is clearly different. Description Length 3.6 mm, width 1.9 mm. Body oval, widest before the middle (fig.262). Head. - Yellow-brown with darkened vertex and front-margin, moderately punctured. Distance be- tween the eyes 1.3 X eye-width. Antennae yellow, first segment yellow-brown, third segment clearly longer than wide, fifth segment slightly longer than fourth (fig. 263). Palpi yellow. Pronotum. - Yellow. Lateral borders straight, finely bordered. Sparsely, basally and anteriorly more densely punctured, on each side of the base 129 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 „eens, Figs. 262-270. Haliplus sharpi (262-267, lectotype; 268-270, from Korea). — 262, dorsal view; 263, antenna; 264, elytral punctuation; 265, dorsal side of hind tibia; 266, prosternal process; 267, lateral view of prosternal process; 268, left paramere; 269, penis; 270, right paramere. 130 with a row of partly widened and darkened punc- tures (fig. 262). Elytra. - Yellow to yellow-brown, rather strong primary puncture-rows, about 36 punctures in the first row. Secondary punctures in the first interval strong and dense, in the other intervals with nor- mal and very small punctures (fig. 264). All punc- tures darkened. Distinct markings along the base, suture and apical point, connected with blotches in the second interval, blotches in the fourth and sixth interval (fig. 262). Ventral side. - Body yellow-red to brown-red, abdominal segments darkened basally, elytral epi- pleura yellow, legs brown-red, darkened towards the coxae. Prosternal process flat, slightly im- pressed in the apical part, densely punctured, slightly narrowed near the coxae, widened towards the apex (fig. 266), lateral plicae complete (fig. 267), anterior edge of prosternum strongly bor- dered, pro-episternum unpunctured. Metasternal process grooved and with a pit in the middle, den- sely punctured, basally as wide as the prosternal process (fig. 266). Coxal plates strongly punctured, width of punctures less than the distance to each other, accompanied by some very small punctures, area along the suture impressed and only weakly and sparsely punctured. Setiferous striole on dorsal side of hind tibia about 4% X the tibia length, longer of the two tibial spurs about % of the length of the first tarsal segment (fig. 265). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs slightly widened, scaly hairs on ventral side. Penis and parameres as in figs. 268-270. Biology Stagnant pools. Caught at light and in a hotel room. Distribution (fig. 310) China, prov. Jiangsu; Japan, Hokkaido and Shi- koku; Korea. Material examined. — 8 ex. - China: 1 9 (lectotype) (MNHN ex coll. Wehncke); 1 ex., Jiangsu, Shanghai, summer 1943, Marist Brothers (ISNB). — Japan: 1 9, Japon, Heller, Dr. Régimbart vidit 1898, © Haliplus sharpi Wehncke, 1880 (paralectotype) (MNHN ex coll. Wehncke); 1 ex., Shikoku, Matsuyama, vii.1959, Light trap, (ZMUC). - Korea: 28, 1 9, Sariwon,12-22.vii.1956, M. Magyar, (ISNB); 1 9, Prov. South Pyongan, Pyongyan, room of Hotel Te-dong, 30.vii.1975, J. Papp and A. Voj- nits (CNCI); 1 @, 1 9, Kyongpuk-do, Yongpung-kun, Hwayon-pond, 2.viii.1988, Sung Hwa Lee (Vondel). 29. Haliplus variegatus Sturm (figs. 271-282) VAN VONDEL: Palaearctic Liaphlus Haliplus variegatus Sturm, 1834: 157. Syntypes probably lost [as most Sturm-types] [not examined]. Haliplus subnubilis Babington, 1836: 177. Syntypes not located [not examined]. Haliplus variegatus var. obsoletus Westhoff, 1881: 43. Syntypes not located [not examined]. Haliplus variegatus ab. pallidior Müller, 1900: 115. Lec- totype 8 (here designated) [Yugoslavia] ‘Castelnuovo, Humler, TYPUS, Museo Civico di Trieste, H.variegatus ab. pallidior M.’ (MSNT) [examined]. Haliplus pantherinus Sahlberg, 1900? (preoccupied). Haliplus leopardinus Sahlberg, 1900: 183 (replacement name for H. pantherinus). Lectotype @ (here desig- nated) [Greece, Kérkira] ‘Corfu, J. Sahlb., Haliplus pantherinus Sahlb. Type, Collect. Hauser, v. leopardi- nus det. Ganglbauer’ (NHMW) [examined] Haliplus transvolgensis Semenov, 1904: 216. Lectotype (here designated) [USSR] ‘Nicolajevskij-Gorodok, Saratov, 431.V.[19]00, Sacharov, Haliplus transvol- gensis mTyp., A. Seminov det. (ZMAS)[examined]. Syn. n. Haliplus variegatus subsp. galilea Sahlberg, 1913: 40. Types not located [not examined]. Haliplus variegatus; Aubé 1836: 28, 1838: 16, Erichson 1837: 184, Schaum 1848:42, Thomson 1854: 485, Ré- gimbart 1877: 354, Westhoff 1881: 43, Seidlitz 1887: 30, Miiller 1900: 114, A pfelbeck 1904: 360, Fiori 1904: 196, Sahlberg 1913: 39, Zimmermann 1920: 317, 1924: 136, Guignot 1928: 145, 1933: 231, 1947: 51, Scholz 1929: 17, F. Burmeister 1939: 214, Balfour- Browne 1940: 152, 1953: 5, Horion 1941: 366, Csiki 1946: 560, Zaitsev 1953: 73, Gueorguiev 1958: 44, 1960: 23, 1981: 401, Lindroth 1960: 46, Lagar Mascaro 1968: 73, Freude 1971: 15, Franciscolo 1972: 60, 1979: 121, Hosseinie 1974: 241, 1978: 167, Ienistea 1974: 208, 1978: 295, Ali 1976: 92, Galewski 1976: 29, Mate- leshko 1977: 68, Fichtner 1981: 328, Holmen 1981: 2, 1987: 116, E. Burmeister 1982: 94, 1987: 184, Leblanc 1984: 8, Vondel 1985: 33, 1989a: 58, Lundberg 1986: 11, Zakharenko & Moroz 1988: 283, Nilsson 1988: 6. Haliplus subnubilis; Schaum 1848: 42, Zimmermann 1920: 317. Haliplus variegatus var. obsoletus; Guignot 1933: 244, Zaitsev 1953: 73, Gueorguiev 1960: 23. Haliplus variegatus ab. pallidior, Zimmermann 1920: 317, Scholz 1929: 17, Guignot 1933: 244, Zaitsev 1953: 74, Gueorguiev 1958: 44, 1960: 23, 1963: 216, Ienistea 1978: 295, Leblanc 1984: 8. Haliplus leopardinus, Apfelbeck 1904: 360, Sahlberg 1913: 39, Zimmermann 1920: 317, Döbler 1976: 182. Haliplus transvolgensts; Zaitsev 1905: 37, Zimmermann 1920: 316 1924: 135, Kinel 1929: 219, Scholz 1929: 18, Burmeister 1939: 214, Csiki 1946: 559. Diagnosis A very variable species in size, colour and mac- ulation. The distance between the eyes, 1.7-2.0 X eye-width, usually distinguishes it from related species of the same size, especially in the Middle- East. Description Length 2.6-3.4 mm, width 1.4-1.8 mm. Body oval, widest before the middle (fig. 271). 131 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 271-279. Haliplus variegutus, Texel, Netherlands. - 271, dorsal view; 272, antenna; 273, elytral punctuation, 274, dorsal side of hind tibia; 275, prosternal process; 276, lateral view of prosternal process; 277, left paramere; 278, penis; 279, right paramere. 132 Head. - Yellow to rust-coloured, darkened ver- tex, rather weakly punctured, on the vertex with stronger darkened punctures, between the eyes with an unpunctured band. Distance between the eyes 1.7-2.0 X eye-width. Antennae yellow to yel- low-brown, third segment longer than the fourth or fifth segment (fig. 272), palpi yellow to rust- coloured. Pronotum. - Yellow to rust-coloured, sometimes anterior and basal margins slightly darkened. Lat- eral borders straight, finely bordered. Sparsely, on the disc almost not, basally strongly punctured, basal punctures darkened (fig. 271). Elytra. - Yellow to rust-coloured, rather strong primary puncture-rows, 26-32 punctures in the first row. Secondary punctures in the first interval fairly strong, in the remaining intervals moder- ately strong, accompanied by very small punctures. All punctures darkened (fig. 273). Extensive mark- ings on the disc, along the suture, on the apex and on the even intervals with extensions to the odd intervals. The central blotches are not always con- nected to the suture (figs. 271, 280-282). Ventral side. - Yellow-red to rust-coloured, ely- tral epipleura yellow, legs yellow-red to rust-co- loured, darkened towards the coxae. Prosternal process flat or slightly convex, narrowed near the coxae (fig. 275), lateral plicae complete (fig. 276), rather coarsely punctured. Metasternal process slightly to strongly and widely grooved, coarsely punctured (fig. 275). Coxal plates sparsely, laterally more densely punctured, with a few very small punctures, apical point broadly rounded. Setiferous striole about ‘4 of the tibia length, with about 5 punctures, longer of the two tibial spurs % the length of the first tarsal segment (fig. 274). Hind tibia about 3 X as long as the first tarsal segment. VAN VONDEL: Palaearctic Liaphlus Figs. 280-282. Haliplus variegatus, elytra. — 280, lectotype of H. variega- tus var. pallidior, 281, from Castelnuovo, Yu- goslavia; 282, from Trenton, England. Male sexual characters. -- First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 277- 279) Biology In stagnant waterbodies with fresh water with often a vegetation of Characeans, Sphagnum or Hypnum, in temporary dune-pools, in peat bogs. The third-stage larva was first described by Schigdte (1864). Distribution (fig. 296) Europe: From Ireland and England to the central and southern part of European USSR, northwards to the south of Norway, Sweden and Finland and southwards to Spain, Italy and Greece. Algeria, Israel, Turkey, Syria. The records from Iraq (Ali 1976), Iran (Hosseinie 1974, 1978) and Afghanis- tan (Gueorguiev 1963) most likely concern other species like abbreviatus or villiersi. The record from West Siberia (Zaitsev 1905, as transvolgen- sis) probably concerns H. maculatus or angust. Remarks Examination of the type of transvolgensis showed that it is conspecific with varzegatzs. In the Mediterranean area small specimens are present, described as pallidior or leopardinus, but in my opinion these are conspecific with variegatus. Material examined. - 425 ex. - Algeria: 2 ex. (BMNH). — Austria: 49 ex. (ZMAN, RMNH, NHMW, RMNH). — Bel- gium: 1 ex. (NHMW). - Denmark: 2 ex. (BMNH, ZMAN). — England: 31 ex.BMNH, ZMAN, RMNH). — France: 69 ex. (MNHN, ZMAN, RMNH, NHMW, Huijbregts, Vondel). - Ger- many: 25 ex. (ZAMAN, RMNH, NHMW, BMNH). - Greece: 1 8 (lectotype leopardinus) (NHMW), 9 ex. (paralectotypes 155 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 283-291. Haliplus villiersi (283-288, holotype; 289-291, from type-locality). - 283, dorsal view; 284, antenna; 285, elytral punctuation; 286, dorsal side of hind tibia; 287, prosternal process; 288, lateral view of prosternal process; 289, left paramere; 290, penis; 291, right paramere. 134 leopardinus) (2 6,3 Q NHMW; 1 9 UZMH; 1 ex. OLML; 1 GB, 1 Q zsMC); 38 ex. (BMNH, ZMAN, NHMW, RMNH, Huij- bregts). - Hungary: 7 ex. (ZMAN, NHMW). — Ireland: 3 ex. (BMNH). - Israel: 2 6, Berekhat Zemorot nr. Ashquelon, 15.1.1981 (NHMW). — Italy: 15 ex. (BMNH, CNCI, NHMW, Huijbregts, Jongema). — Netherlands: 108 ex (RMNH, Schilthuizen, Cuppen, ZMAN, BMNH). — Poland: 6 ex. (BMNH, ZMAN). — Spain: 4 ex. (ZMAN, NHMW). - Sweden: 2 ex. (BMNH). — Switzerland: 1 ex. (BMNH). — Turkey: 1 Q (MNHN). — USSR: 1 9, (lectotype transvolgensis) (ZMAS). — Yugoslavia: 8 ex. (lectotype and 4 paralecto- types of pallidior) (MSNT); 10 ex. (NHMW, CNCI). — Local- ity not traced or unknown: 30 ex. (BMNH, ZMAN, MHNG, RMNH, NHMW). 30. Haliplus villiersi Legros (figs. 283-291) Haliplus villiersi Legros, 1972: 573: Holotype @, ‘Iran, Zabol, IV.1965, Museum Paris, Mission Franco-Ira- ‘ nienne 1965, A, Type, Haliplus (Liaphlus) villiersi n. sp., C. Legros det.’ (MNHN) [examined]. Diagnosis A variable species, that is closely related to ab- breviatus and variegatus, from which it can be distinguished by the usually impressed prosternal process. Maybe abbreviatus and villiersi are con- specific. Description Length 3.0-3.4 mm, width 1.7-2.0 mm. Body oval, sides subparallel, shoulders a little pro- nounced (fig. 283). Head. - Yellow-brown, weakly punctured. Dis- tance between the eyes about 1.6 X eye-width. Antennae (fig. 284) and palpi yellow.. Pronotum. - Yellow-red, lateral borders straight, narrowed anteriorly, finely bordered. Weakly, along the base more strongly, on the disc more sparsely punctured, basal punctures slightly dar- kened (fig. 283). Elytra. - Yellow to yellow-brown, rather weak to moderately strong primary puncture-rows, 30 to 35 punctures in the first row. Punctures of 8th and 9th row with long hairs. Secondary punctures in two types: moderately strong or very weak (fig. 285). All punctures darkened. Usually brown markings on the suture, apex, even intervals and sometimes in the first interval (fig. 283), the markings can be very weak. Ventral side. - Body yellow to yellow-red, elytral epipleura yellow, legs yellow-red. Prosternal pro- cess flat or slightly impressed apically, coarsely punctured, slightly narrowed near the coxae (fig. 287), lateral plicae complete (fig. 288). Metasternal process flat or grooved in the middle and with a central pit (fig. 287), weakly punctured. Setiferous VAN VONDEL: Palaearctic Liaphlus striole on dorsal side of hind tibial about one third of the tibia length, longer of the two tibial spurs % the length of the first tarsal segment (fig.286). Male sexual characters. - First three tarsal seg- ments of fore- and midlegs widened, scaly hairs on ventral side. Penis and parameres as in figs. 289- 291. Distribution (fig. 305) Iran. Material examined. - 6 ex. - Iran: 1 & (holotype); 1 6, 4 ©, Zabol, 19.iv.1965, Mission Franco-Iranienne 1965 (1 Q allotype) (MNHN) Unnamed material The specimens I failed to determine are listed below. Some of them might belong to still undes- cribed species. Species near H. diruptus. - [China] ‘Yunnan Fou, Yunnan, R. Mouchamps det. 63, Haliplus di- ruptus B.Br. (1 9, 1 @ without aedeagus) (ISNB). Belonging to Haliplus diruptus or H. davidi or to an undescribed species. - [China, Beijing] ‘Peping, Collector C. F. Wu, Col. Gschwendtner, simplex Clark Gschwendtner’ (2Q)(OLML). Possibly be- longing to H. diruptus or H. davidi. - [China] ‘Yunnan Fou, Yunnan, H. Perrot; ® (MNHN). Pos- sibly belonging to H. diruptus or H. davidi. Species near H. variegatus. - [USSR] ‘Ross. O. (2 92) (BMNH). Possibly belonging to an undes- cribed species. Species near H. ortali. - [Iran] ‘Perse, Susa, iv.1899, Escalera, Haliplus sp.x., Dr. Régimbart vidit 1900 (1 2) (MNHN). Possibly belonging to H. ortali. ACKNOWLEDGEMENTS I am very grateful to Dr. R. Angus (Englefield Green) for his valuable help and for critically read- ing the manuscript. I wish to express my sincere thanks to the fol- lowing persons for placing material or information at my disposal: Dr. G. Alberti (Trieste, MSNT), Dr. L. Baert (Brussels, ISNB), Dr. A. L. van Berge Hene- gouwen (The Hague, MOG), Dr. O. Bistròm (Hel- sinki, UZMH), Dr. M. Brancucci (Basel, NHMB), Dr. M. J. D. Brendell (London, BMNH), mr. B. Brugge (Amsterdam, ZMAN), Dr. J. G. M. Cuppen (Ede), Dr. R. Danielsson (Lund, MZLU), Dr. K. Desender (Brussels, ISNB), mr. M. B. P. Drost (Wadenoijen), Dr. J. P. Duffels (Amsterdam, ZMAN), Dr. Th. G. Giessen (Gaanderen), Dr. F. Gusenleitner (Linz, OLML), Dr. F. Hieke (Berlin, ZMHB), mr. M. Hiel- kema (Gouda), Dr. S. J. Hine (London, BMNH), mr. 135 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 M. Holmen (Copenhagen, ZMUC), Dr. J. Huijbregts (Leiden, RMNH), Dr. M. Jäch (Vienna, NHMW), mr. Y. Jongema (Wageningen), Dr. B. Korotyaev (Le- ningrad, ZMAS), Dr. J. Krikken (Leiden, RMNH), Dr. P. Leblanc (Piney), Dr. Sung Hwa Lee (Taegu), Dr. L. LeSage (Ottawa, cncı), Dr. P. Lindskog (Stock- holm, NHRS), Dr. I. Löbl (Genève, MHNG), Dr. K. Majer (Brno, MMBC), Dr. E. Matsui (Hondo City), Dr. T. Nakane (Chiba), Dr. R. Ortal (Jerusalem, HUJI), mrs. H. Perrin (Paris, MNHN), Dr. R. Poggi (Genua, MCSN), mr. EF F. Repko (Leersum, RIN), Dr. G. Scherer (Munich, zsMc), Dr. M. Schilthuizen (Leiden), Dr. A. Smetana (Ottawa, CNCI), mr. G. Stobbe (Leeuwarden, FNML), Dr. R. zur Strassen (Frankfurt am Main, sMFD), mr. H. Vallenduuk (Boxtel). The Uyttenboogaart-Eliasen Foundation is acknowledged for financial support. REFERENCES Alfieri, A., 1976. The Coleoptera of Egypt. - Mémoires de la Société Entomologique d’Egypte 5: 38. Ali, H. A., 1976. Preliminary study on the aquatic beetles of Iraq (Haliplidae, Coleoptera). — Bulletin of the Basrah Natural History Museum 3: 89-94. Apfelbeck, V., 1904. Die Kaferfauna der Balkanhalbinsel mit Beriicksichtigung Klein-Asiens und der Insel Kreta. I: Familienreihe Caraboidea. — Friedländer, Berlin, IX+422 pp. Arnett, R. H., & G. A. Samuelson, 1986. The Insect and Spider Collections of the World. — E. J. Brill, Gaines- ville, 220 pp. Aubé, C., 1836. Hydrocanthares. - In: Dejean, Boisduval & Aubé. Iconographie et Histoire Naturelle des Co- léoptères d'Europe V. Méquignon-Marvis, Paris, 415 PP. Aubé, C., 1838. Species général des Hydrocanthares et Gyriniens, 6. - Méquignon-Marvis, Paris, XVI+804 PP. Babington, C. C., 1836. Observations on Haliplus ferru- gineus of Authors, being an attempt at its Subdivision into several Species. - Transactions of the Entomolog- ical Society of London 1: 175-179, 1 pl. Balfour-Browne, F., 1936. Systematic notes upon British aquatic Coleoptera. Part XI. Haliplidae. - Entomolo- gist's Monthly Magazine 72: 68-77. Balfour-Browne, F., 1938. Systematic Notes upon British Aquatic Coleoptera, I, Hydradephaga. — Nathaniel Lloyd, London, 95 pp. Balfour-Browne, F, 1940. Brit- ish Water Beetles, I. - Ray Society, London, XIX+375 Pps pl. Balfour-Browne, F., 1951. The aquatic Coleoptera of Ire- land. - Entomologist’s Gazette 2: 1-52. Balfour-Browne, F, 1953. Coleoptera Hydradephaga. - Handbooks for the identification of British Insects 4 (3)1833: Balfour-Browne, J., 1946. The Aquatic Coleoptera of Manchuria (Weymarn Collection). - Annals and Mag- azine of Natural History (11), 13: 433-460. Balfour- Browne, J., 1948. The aquatic coleoptera of New- foundland & Nova Scotia. - The Canadian Entomol- ogist 80: 156-165. 136 Balfour-Browne, J., 1951. Coleoptera: Haliplidae, Dytis- cidae, Gyrinidae, Hydraenidae, Hydrophilidae. - Ex- pedition to South-West Arabia 1937-8, I, 16: 179-219. Belyashevskiy, N. N., 1989. Predaceous water beetles (Coleoptera, Hydradephaga) of the Slovechansk-Ov- ruch ridge and adjacent regions of the Poles’e lowland. — Entomological Review 68(3): 122-137. Bertrand, H., 1928. Les variétés pyrénéennes de l’Haliplus fulvus F. (Col. Haliplidae). - Bulletin de la Société Entomologique de France : 87-90. Bertrand, H., 1949. Récoltes de Coléopteres aquatiques (Hydrocanthares) dans les Pyrénées; observations écologiques. - Bulletin de la Société Entomologique de France 74: 24-38. Beutel, R. G. & S. Ruhnau, 1990. Phylogenetic Analysis of the Genera of Haliplidae (Coleoptera) Based on Characters of Adults. - Aquatic Insects 12: 1-17. Blatchley, W. S., 1910. An illustrated descriptive catalogue of the coleoptera or beetles known to occur in Indiana. -The Nature Publishing Co., Indianapolis. Brinck, P., 1942. Die von J. W. Zetterstedt und C. G. Thomson neu beschriebene Halipliden und Dytis- ciden. — Kungliga Fysiografiska Sällskapets i Lund Förhandlingar Bd. 12 Nr. 11 :1-17. Burmeister, E. G., 1982. Ein Beitrag zur aquatischen Ka- ferfauna von Walcheren, Niederlande (Insecta, Co- leoptera) unter besonderer Berücksichtigung von ha- lophilen und halobionten Arten. — Zoologische Bijdragen 28: 85-99. Burmeister, E. G., 1985. Interessante Wasserkäferarten einer Aufsammlung aus Israel und dem Sinai. Ana- caena jordanensis sp. n. (Coleoptera, aquatica). - En- tomofauna 6: 37-56. Burmeister, E. G., K. Dettner & M. Holmen, 1987. Die Hydradephaga Sardiniens. — Spixiana 10: 157-185. Burmeister, F., 1939. Biologie, Ökologie und Verbreitung der Europäischen Käfer auf systematischer Grund- lage, I, Adephaga. - Goecke & Evers, Krefeld, 307 pp. Cho, P. S., 1969. Insecta II. — Illustrated Encyclopedia of Fauna & Flora of Korea 10: 1-970. Clairville, J. de, 1806. Entomologie Helvétique II. - Orell, Fussli & Compagnia, Zürich, 290 pp, 16 tab. . Clark, H., 1863. Descriptions of new East-Asiatic Species of Haliplidae and Hydroporidae. - Transactions of the Entomological Society of London 3: 417-428. Crotch, G. R., 1873. Revision of the Dytiscidae of the United States. - Transactions of the American Ento- mological Society 4: 383-424. Csiki, E., 1946. Die Käferfauna des Karpaten-Beckens, I, Algemeiner Teil und Caraboidea. - Budapest, 796 pp. Delarouzee, C., 1857. Bulletin entomologique 3e Trimes- tre 1857. -Annales de la Société Entomologique de France 5: XCIV-XCV. Dobler, H., 1976., Katalog der in den Sammlungen des ehemaligen Deutschen Entomologischen Institutes aufbewahrten Typen XIV (Coleoptera: Dytiscidae, Haliplidae, Gyrinidae, Rhysodidae, Paussidae). — Bei- träge zur Entomologie 26: 179-186. Erichson, W. F, 1837. Die Käfer der Mark Brandenburg I, 1. - Morin, Berlin, 384 pp. Escalera, M. M. de la, 1914. Los Coleopteros de Marru- ecos. — Trabajos Museo Nacional de Ciencias Natu- rales 11: 56-57. Fabricius, J. C., 1787. Mantissa Insectorum, I. - Hafniae. Fabricius, J. C., 1801. Systema Eleutheratorum I. - Kiel, XXIV+506 pp. Falkenström, G., 1932. Vorläufige Mitteilung über die neuen Halipliden und Dytisciden, von Dr. D. Hum- mel in den Jahren 1927-30 während Dr. Sven Hedins China-Expedition eingesammelt. — Entomologisk Tidskrift 53: 191-192, 1 pl. Falkenström, G., 1933. Schwedisch-chinesische wissen- schaftliche Expedition nach den nordwestlichen Pro- vinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sü Ping-chang. Insekten, 15, Coleoptera. 1. Ha- liplidae und Dytiscidae. - Arkiv för Zoologi 27A: 1- IR Falkenström, G., 1936. Halipliden, Dytisciden und Gyri- niden aus West- und Central-China (Coleoptera). - Lingnan Science Journal 15: 79. Falkenström, G., 1941. Stockholmstraktens Vatten-Co- leoptera. - Entomologisk Tidskrift 61: 88-92. Fichtner, E., 1981. Beiträge zur Insektenfauna der DDR: Coleoptera-Haliplidae. - Beiträge zur Entomologie 31 (2): 319-329. Fiori, A., 1904. Studio critico dei Dyticidi Italiani. - Riv- ista Coleotterologica Italiana 2: 186-205. Fiori, A., 1914. Appunti sulla fauna coleotterologica del- l'Italia meridionale e della Sicilia. - Rivista Coleotte- rologica Italiana 12: 166. Foster, G. N., 1981. Atlas of British Water Beetles- Part 1. - Balfour-Browne Club Newsletter 22: 1-18. Franciscolo, M. E., 1972. Hydrodephaga of Yugoslav Ad- riatic Islands, I. - Acta Entomologica Jugoslavica 8: 55- 90. Franciscolo, M. E., 1979. Coleoptera. Haliplidae, Hygro- biidae, Gyrinidae, Dytiscidae. - Fauna d'Italia 14: 804 PP. Freude, H., 1971. Familie: Haliplidae, Wassertreter. In: H. Freude, K. W. Harde & G. A. Lohse. Die Käfer Mitteleuropas 3: 8-15. Galewski, K., 1976. Chrzaszcze Coleoptera, Flisakowate -Haliplidae, Hygrobiidae. - Klucze do Oznaczanıa Owadéw Polski 19(5-6): 13-52. Gislason, G. M., 1977. Islenskar vatnabjöllur. - Natturu- fraedingurinn 47: 154-159. Gschwendtner, L., 1921. Beiträge zur Kenntnis und Bes- timmung paläarktischer Schwimmkäfer (Haliplidae et Dytiscidae). - Entomologischer Anzeiger 1: 56-63. Guéorguiev, V. B., 1958. Verzeichnis der Wasserkäfer Bulgariens. -Entomologische Blätter (für Biologie und Systematik der Käfer) 54: 44-51. Guéorguiev, V. B., 1960. Contribution a la connaissance des Coleopteres Hydrocanthares de Yougoslavie. Acta Musei Macedonici Scientiarum Naturalium 7: 19-39. Guéorguiev, V. B., 1963. Contribution a l'étude des Co- léoptéres Hydrocanthares (Haliplidae et Dytiscidae) d'Afghanistan. - Opuscula Entomologica 28: 215-222. Guéorguiev, V. B., 1981. Résultat de l'expedition zoolo- gique du musée national de Prague en Turquie, Co- leoptera: Haliplidae, Dytiscidae, Gyrinidae. - Acta En- tomologica Musei Nationalis Pragae 40: 399-424. Guignot, F, 1928. Notes sur les Haliplus du groupe fulvus F. - Annales de la Société Entomologique de France RSA Guignot, F., 1930. Notes sur quelques Haliplus (Col. Ha- liplidae). - Bulletin de la Société Entomologique de France: 71-75. Guignot, F., 1933. Les Hydrocanthares de France. - Dou- ladoure, Toulouse, XV+1057 pp. Guignot, F, 1947. Coléoptères Hydrocanthares. - Faune de France 48: 1-286. Guignot, F, 1950. Trente-deuxieme note sur les Hydro- VAN VONDEL: Palaearctic Liaphlus canthares. -Revue Française d’Entomologie 17: 97- 104. Guignot, F., 1954. Entomological results from the Swed- ish expedition 1934 to Burma and British India. Co- leoptera: Haliplidae & Dytiscidae. - Arkiv for Zoologi 6 (32): 563-567. Guignot, F., 1955. Sur la Systématique des Haliplus (Col. Haliplidae). - Mémoires de la Société Royale d’Ento- mologie de Belgique 27: 289-296. Guignot, F., 1959. Revision des Hydrocanthares d'Afrique (Coleoptera Dytiscoidea). - Annales du Musée Royal du Congo Belge. Sér. 8, 70 (1): 1-165. Gundersen, R. W. & C. Otremba, 1988. Haliplidae of Minnesota. — Scientific Publications of the Science Museum of Minnesota. New series 6(3): 1-43. Gyllenhal, L., 1808. Insecta Suecica, Coleoptera I. - Leve- rentz, Scaris, XII+-572 pp. Hatch, M. H., 1953. The beetles of the Pacific Northwest. Part I. Introduction and Adephaga. - University of Washington Publications in Biology 16: 1-340. Hilsenhoff, W. L. & W. U. Brigham, 1978. Crawling water beetles of Wisconsin (Coleoptera: Haliplidae). - The Great Lake Entomologist 11: 11-22. Holmen, M., 1981. A survey of Denmark's Haliplidae (Coleoptera) with remarks on zoogeography and autecology. - Entomologiske Meddelelser 49: 1-14. Holmen, M., 1987. The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark, I, Gyrinidae, Haliplidae, Hygrobiidae and Noteridae. - Fauna Entomologica Scandinavica 20: 1-168, 1 pl. Horion, H., 1941. Faunistik der Deutschen Kafer, I, Ade- phaga-Caraboidea. - Diisseldorf, 464 pp. Hosseinie, S. O., 1974. Water Beetles found in the vicinity of Shiraz, Iran (Coleoptera: Dytiscidae, Noteridae, Haliplidae, Gyrinidae and Hydrophilidae). - Coleop- terist’s Bulletin 28(4): 237-243. Hosseinie, S. O., 1978. Aquatic Coleoptera from Southw- estern Iran (Haliplidae, Dytiscidae and Gyrinidae with a note on Hydrophilidae). - Coleopterist’s Bul- letin 32(2): 167-175. Ienistea, M. A., 1974. Beitrag zur Kenntnis der Hali- pliden Rumäniens (Coleoptera). - Travaux du Mu- séum d'Histoire Naturelle "Gr. Antipa” 15: 203-211. Ienistea, M. A., 1978. Hydradephaga und Palipicornia. — In: J. Illies (ed. ), Limnofauna Europaea. Stuttgart. Jackson, D. J., 1952. Observations on the capacity for flight of water beetles. - Proceedings of the Royal Entomological Society of London 27: 57-70. Jackson, D. J., 1956. The capacity for flight of certain water beetles and its bearing on their origin in the Western Scottish Isles. - Proceedings of the Linnean Society of London 167: 76-96. Jackson, D. J., 1973. The influence of flight capacity on the distribution of aquatic Coleoptera in Fife and Kinross- shire. - Entomologist’s Gazette 24: 247-293. Kano, T., & K. Kamiya, 1931. Two new species of Hali- plidae from Japan. Transactions of the Kansai Ento- mological Society 2: 1-4, 1 pl. Kinel, J., 1929. Einige Bemerkungen zur Bestimmungsta- belle. 97 Heft, Haliplidae (Col. ). - Polskie Pismo Entomologiczne 8: 217-220. Kolossov, I., 1931. Synonymische und systematische Be- merkungen über Coleopteren palaearcticis. - Entomo- logische Blätter (für Biologie und Systematik der Käfer) 27: 116-118. Konev, A. A., 1976. Water beetles of the suborder Ade- 157 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 phaga (Coleoptera) in Central Kazakhstan. — Ento- mological Review 55: 56-58. Lagar Mascaro, A. L., 1968. Los Haliplidae (Coleoptera Dytiscoidea) de Cataluna. - Miscellanea zoologica II (3): 65-73. Larson, D. J., 1987. Aquatic coleoptera of peatlands and marshes in Canada. In: D. M. Rosenberg & H. V. Danks. Aquatic insects of peatlands and marshes in Canada. - Memoirs of the Entomological Society of Canada 140: 99-132. Leblanc, P., 1984. Iconographie d’Hydrocanthares Cham- penois (suite) ‘Col. Haliplidae’. - Bulletin d’Entomol- ogie Champenoise 3: 8, 40. Legros, C., 1972. Contribution a la faune d’Iran. 23. Co- leopteres hydrocanthares. - Annales de la Société En- tomologique de France 8 (3): 573-578. Lindroth, C. H. (ed. ), 1960. Catalogus Coleopterorum Fennoscandiae et Daniae. - Entomoliska Sallskapet, Lund, 476 pp. Linnaeus, C., 1767. Systema Naturae, ed. 12, II. - Laur. Salvii, Holmiae, 553-1327. Lundberg, S., 1986. Catalogus Coleopterorum Sueciae. - Entomologiska Foreningen och Naturhistoriska Riks- museet, Stockholm, 302 pp. Marseul, S. A. de, 1882. Nouveau repertoire contenant les descriptions des espèces de Coléoptères de l'Ancien monde &c. . - L'Abeille Journal d’Entomologie 20: 105-112. Marsham, T., 1802. Entomologia Brittannica I, Coleop- tera. - London, XX XI+548 pp. Mateleshko, M. F, 1977. Water beetles and their distri- bution in water bodies of the Transcarpathian Region. - Vestnik Zoologii 10: 67-73. Minoransky, V. A.& N. B. Dzumailo, 1974. On fauna of water beetles in the Rostov Region. - Vestnik Zoolo- gii 7: 25-32. Motschulsky, V. de, 1860. Coléoptères de la Sibérie orien- tale et en particulier des rives de l'Amour. - Schrenck's Amur-Reise II, Coleopteren: 79-257. Miiller, J., 1900. Haliplidae, Hygrobiidae, Dytiscidae et Gyrinidae Dalmatiae. - Verhandlungen der Zoolo- gisch-Botanischen Gesellschaft in Wien 50: 112-121. Müller, J., 1926. Coleopterologische Notizen X. - Wiener Entomologische Zeitung 43: 167-171. Munster, T., 1922. Om nogle Haliplider. - Videnskaps- selskapets Skrifter. I. Mat. Naturv. Klasse 9: 3-15. Nakane, T., 1963. Iconographia Insectorum Japonicorum, II, Coleoptera. - Hokuryukan, Tokyo, 413 pp. Nakane, T., 1985a. A revisional study on Japanese species of the genus Haliplus Latreille (Coleoptera, Halipli- dae). - Kita-Kyüshü no Konchü 32: 61-67. Nakane, T., 1985b. [Japanese title]. - Gekkan-Mushi 171: 40-41. Nakane, T., 1987. The beetles of Japan (new series) 80: 26-30. Nilsson, A., 1988. Laccophilus obsoletus is a Haliplus. - Balfour-Browne Club Newsletter 43: 6. Ohkura, M., 1962. Haliplidae. - In: T. Nakane (ed. ), Coloured illustrations of The Insects of Japan. Coleop- tera. - Hoikusha, Osaka. Pearce, E. J., 1939. New and additional Haliplid records for Eire and some notes on variation in British spec- imens of Haliplus fulvus F. (Coleopt. ). - Entomolo- gist's Monthly Magazine 75: 32-36. Penecke, K. A., 1901. Coleopterologische Miscellen II. - Wiener Entomologische Zeitung 20: 11. Perris, M. E., 1857. Nouvelles excursions dans les grandes 138 Landes. - Annales de la Société Linnéenne de Lyon 4: 83-180. Redtenbacher, W., 1842. Quaedam genera et species Co- leopterorum Archiducatus Austriae nondum descrip- torum. - Vindobonae, Ueberreuter, 31 pp. Regimbart, M., 1877. Enumeration des Dytiscides et Gyr- inides receuillis par Ch. Piochard de la Brulerie dans ses voyages en Orient. - Annales de la Société Ento- mologique de France 15 (7): 347-354. Régimbart, M., 1886. Dytiscidae et Gyrinidae nouveaux de la collection du musée de Leyde. - Notes from the Leyden Museum 8: 139-144. Régimbart, M., 1899. Révision des Dytiscidae de la Ré- gion Indo-Sino-Malaise. - Annales de la Société Ento- mologique de France 68: 186-. Régimbart, M., 1901. Notes sur quelques Dytiscides d’Eu- rope (Col.). — Bulletin de la Société Entomologique de France 70: 323-325. Riha, P., 1955. Ergebnisse der Tschechoslovakischen Is- land-expedition 1948 (Col., Haliplidae, Dytiscidae). - Acta Entomologica Musei Nationalis Pragae 30: 55- 60. Roberts, C. H., 1913. Critical notes on the species of Haliplidae of America north of Mexico with descrip- tions of new species. -Journal of the New York Ento- mological Society 21: 91-123. Ruette, R. de, 1970. A catalogue of types of Coleoptera in the Canadian National Collection of Insects. — Me- moirs of the Entomological Society of Canada 72: 1- 134. Sahlberg, J., 1900. Coleoptera mediterranea et rosso-asi- atica. - Ofversigt af Finska Vetenskaps-Societetens Forhandlingar 42: 174-208. Sahlberg, Jey LOW Coleoptera mediterranea et rosso-asi- atica nova et minus cognita. - Ofversigt af Finska vetenskaps-Societetens Förhandlingar 50: 1-94. Sahlberg, J., 1913. Coleoptera mediterranea orientalia. — Öfversigt af Finska Vetenskaps-Societetens Förhan- dlingar 55: 1-268. Sato, M., 1960. Aquatic Coleoptera from Anami-Oshima of the Ryukyu Islands (I). - Kontyu 28: 251-254. Sato, M., 1984. Taxonomic notes on the aquatic Coleop- tera of Japan, I. - Coleopterists’ News 65: 1-4. Schaller, J. G., 1783. Neue Insekten. - Abhandlungen der Naturforschenden Gesellschaft zu Halle: 217-332. Schaum, H., 1848. Bemerkungen über brittische Lauf- und Wasserkafer. - Entomologische Zeitung. Stettin 9: 34-44. Schaum, H., 1859. Beiträge zur europäischen Käferfauna. - Berliner Entomologischer Zeitschrift 3: 42-59. Schaum, H., 1864. Die egyptischen Dytisciden. Berliner Entomologischer Zeitschrift 8: 105-108. Schigdte, J. C., 1864. De metamorphosi Eleutheratorum observationes. - Naturhistorisk Tidsskrift 3 (3): 131- 224. Scholz, R., 1915. Beitrag zur Kenntnis und Verbreitung europäischer Wasserkäfer (Haliplidae, Dytiscidae). — Entomologische Blätter (für Biologie und Systematik der Käfer) 11: 232-250. Scholz, R., 1916. Wissenschaftl. Ergebnisse der Bearbei- tung von O. Leonhard's Sammlungen. - Entomolo- gische Mitteilungen 5: 163-182. Scholz, R., 1929. Haliplidae (Col. ). - Bestimmungs-Ta- bellen der europàischen Coleopteren. 97: 1-19. Seeger, W., 1971a. Morphologie, Bionomie und Ethologie von Halipliden, unter besonderer Berücksichtigung funktions-morphologischer Gesichtspunkte (Halipli- dae; Coleoptera). - Archiv für Hydrobiologie 68: 400- 435. Seeger, W., 1971b. Autökologische Laboruntersuchungen an Halipliden mit zoogeographischen Anmerkungen (Haliplidae; Coleoptera). - Archiv fiir Hydrobiologie 68: 528-574. Seeger, W., 1971c. Die Biotopwahl bei Halipliden, zu- gleich ein Beitrag zum Problem der syntopischen (Sympatrischen s. str. ) Arten (Haliplidae; Coleop- tera). - Archiv fiir Hydrobiologie 69: 155-199. Seidlitz, G., 1887. Bestimmungs-Tabellen der Dytiscidae und Gyrinidae des Europäischen Faunengebietes. — Verhandlungen des Naturforschenden Vereins in Briinn 25: 1-136. Semenov, A., 1904. Analecta coleopterologica. - Entomo- logicheskoe Obozrénie 4: 201-202. Sharp, D., 1884. XXI. The water-beetles of Japan. - Transactions of the Entomological Society of London 32: 439-464. Stephens, J. F., 1829. Illustrations of British Entomology. Mandibulata II. Baldwin & Cradock, London, 200 pp., 5 pl. Strand, A., 1944. Nord-Norges Coleoptera. - Tromso Museums Aarshefter 67: 1-626. Sturm, J., 1834. Deutschlands Insekten, 8, Kafer. - Nürn- berg, 2+170 pp., 18 pl. Takizawa, M., 1931. The Haliplidae of Japan. - Insecta Matsumarana 5: 137-143. Thomson, C. G., 1854. Öfversigt af der arter inom fa- miljen Dytisci, som blifvit anträffade pa Skandina- viska halfön. - Svenska Vet. Akad. Handl. : 177-237. Vondel, B. J. van, 1985. Verspreiding van Haliplidae in Nederland. Een tussenstand. - Nieuwsbrief European Invertebrate Survey Nederland 16: 31-34. Vondel, B. J. van, 1986. Description of the second and third-instar larvae of Haliplus laminatus (Schaller) with notes on the subgeneric status (Coleoptera: Ha- liplidae). - Entomologische Berichten, Amsterdam 46: 128-132. Vondel, B. J. van, 1988. Description of Haliplus kulleri n. sp. from Israel (Coleoptera: Haliplidae). - Entomolo- gische Berichten, Amsterdam 48: 97-98. Vondel, B. J. van, 1989a. Watertreders in de Achterhoek en de Liemers. - Natuur en Landschap in Achterhoek en Liemers 3: 54-60. Vondel, B. J. van, 1989b. Water beetling in Wales. — Balfour-Browne Club Newsletter 45: 17-18. Wallis, J. B., 1933. Revision of the north American species (north of Mexico), of the genus Haliplus Latreille. - Transactions of the Royal Canadian Institute 19(1): 1- 76. VAN VONDEL: Palaearctic Liaphlus Wallis, J. B., 1973. An annotated list of the Hydrodephaga (Coleoptera: Insecta) of Manitoba and Minnesota. — Questiones Entomologicae 9: 99-114. Wehncke, E., 1874. Fünf neue europäische Dytisciden. — Berliner Entomologische Zeitschrift 16: 135-136. Wehncke, E., 1880. Neue Haliplus. - Stettiner Entomo- logische Zeitung 41: 72-75. Wehncke, E., 1883. Neue Halipliden. - Deutsche Ento- mologische Zeitschrift 27: 145-146. Westhoff, F., 1881. Die Kafer Westfalens I. - Max Cohen, Bonn, XXVIII+140 pp. White, C. E., F. N. Young & N. M. Downie, 1985. A Checklist of the Aquatic Coleoptera of Indiana. - In- diana Academy of Science 94: 357-369. Wu, C. F, 1932. Aquatic Insects of China. Article XI. Catalogue of Chinese Haliplidae. - Peking Natural History Bulletin 7: 339-343. Yoon, I. B., 1988. Aquatic Insects. - Illustrated Encyclope- dia of Fauna & Flora of Korea 30: 1-870. Yoshimura, S., 1932. On a new species of Haliplus from Japan. - Mushi 5: 102-103. Zaitsev, P., 1905. Notizen über Wasserkäfer (Coleoptera aquatica). - Revue Russe d’Entomologie 5: 37-41. Zaitsev, P., 1907. Berichtigungen und Zusätze zu den Haliplidae, Dytiscidae und Gyrinidae in den neuesten Katalogen der Coleopteren. - Entomologicheskoe Ob- ozrenie 7: 114-124. Zaitsev, F. A., 1953. Coleoptera. Amphizoidae, Hygrobii- dae, Haliplidae, Dytiscidae, Gyrinidae. - Fauna of the U. S. S. R. 58 (4): 1-401. Zakharenko, V. B. & M. D. Moroz, 1988. Material on fauna of water beetles (Coleoptera: Haliplidae, Dytis- cidae, Gyrinidae) of Byelorussia. - Entomologicheskoe Obozrénie 67: 282-290. Zetterstedt, J. W., 1828. Fauna Insectorum Lapponica I. -Schulz, Hammone, 20+563 pp. Zimmermann, A., 1920. Dytiscidae, Haliplidae, Hygro- biidae, Amphizoidae. - Coleopterorum Catalogus 71: 15325. Zimmermann, A., 1924. Die Halipliden der Welt. - En- tomologische Blätter (für Biologie und Systematik der Käfer) 20 (1): 1-16; (2): 65-80; (3): 129-144; (4): 193- 213. Zimmermann, A., 1927. Fauna sumatrensis. Beitrag Nr. 45. Revision der Haliplidae et Dytiscidae von Suma- tra. - Supplementa Entomologica 16: 1-44. Zimsen, E., 1964. The type material of I. C. Fabricius. — Munksgaard, Copenhagen, 654 pp. Received: 10 October 1990 Accepted: 3 December 1990 139 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Fig. 292. Distribution of Haliplus fulvus (dots: material examined, stars: literature records). 140 VAN VONDEL: Palaearctic Liaphlus “(sp102a1 3IMILI9I] :SILIS ‘paurwexa jerJorew !SIOP) 5717801490 ‘H ‘967 ‘(Sp10391 2IMIEIONT :SIEIS pouruexa [eIJaIVU :sIOP) S177098ar7[ ‘H ‘667 ‘(SP109931 a1MeI9N] :SsIeIS ‘PIUTWEXI JEIISIEUI !SIOP) 5772404944 ‘H ‘467 ‘(SP10991 3IMEI9IN] :SIVIS paurwexa jerorew :sIOP) sagrunur ‘H ‘€67 “dds s#]dirH jo uonnqinsiG ‘967-€67 ‘SSA 141 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 297-299. Distribution of Haliplus spp. 297, H. guttatus (dots: material examined, stars: literature records); 298, H. andalusicus (dots: material examined, stars: literature records); 299, H. rubidus (dots: material examined, stars: literature records). 142 VAN VONDEL: Palaearctic Liaphlus Figs. 300-306. Distribution of Haliplus spp. 300, H. maculatus (dots: material examined, stars: literature records); 301, H. dalmatinus (dots: material examined); 302, H. jaechi (dot) and H. kulleri (star: material examined); 303, H. gafnyi (stars) and H. ortali (dots); 304, H. angusi (dot) and H. astrakanus (star); 305, H. abbreviatus (dots: material examined, stars: literature records) and H. villiersi (cross: material examined); 306, H. maculipennis (dot: material examined). 143 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 307-312. Distribution of Haliplus spp. 307, H. pulchellus (dots: material examined); 308, H. eximis (dots: material examined, stars: literature records); 309, H. chinensts (dots: material examined) and H. ovalis (cross: material examined); 310, H. kotoshonis (dot: material examined, star: literature records) and H. sharpi (cross: material examined); 311, H. diruptus (dots: material examined, stars: literature records) and H. basinotatus (cross: material examined, square: literature records); 312, H. excoffieri (star), H. davidi (dot) and H. holmeni (cross). 144 Tijdschrift voor Entomologie Volume 134, no. | Articles | C. Gielis Pterophorus Schaffer in the Ethiopian region (Lepidoptera: Pterophoridae: Pterophorinae): new species and checklist. 9 R. V. Hensen Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae). 31 P. Huemer Caryocolum delphinatella (Constant) sp. rev., a senior synonym of C. fiorii (Klimesch) (Lepidoptera: Gelechiidae). 35 I. Lansbury Cuticular blades and other structures of Diaprepocoris Kirkaldy and Stenocorixa Horvath (Heteroptera: Corixidae). 47 N. Nieser & P. Chen Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), |. 69 R. Puplesis, S. Seksjaeva & V. Sruoga Leaf-mining Lepidoptera (Nepticulidae, Bucculatricidae, Gracillariidae) from Ulmus in northern Caspiya (Kaspia). 75 B.J. van Vondel Revision of the palaearctic species of Haliplus subgenus Liaphlus Guignot (Coleoptera: Haliplidae). Book reviews 68 Medvedev, G. S. (ed), 1990. Keys to the Insects of the European Part of the USSR. Volume IV. Lepidoptera, Part 2. English Translation. [E. J. van Nieukerken]. 74 Emmet, A. M., 1991. The scientific names of the British Lepidoptera. Their history and meaning. [E. J. van Nieukerken].; Rubtsov, |. A., 1990. Blackflies (Simuliidae). - Fauna of the USSR. Diptera, Vol. 6, Part 6. [J. van Tol]. © Nederlandse Entomologische Vereniging, Amsterdam Published | July 1991 ISSN 0040-7496 Le 2 ver Volume 134, no. 2, 1991 a): NEEN 0040-7494 Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Published by the Nederlandse Entomologische Vereniging o Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zoölogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, ’s-Gravenhage CEPHALACHORUTES GEN. N., A NEW GENUS MCZ LIBRARY JAN 21 1992 A. BEDOS & L. DEHARVENG Université Paul Sabatier, Toulouse, France HARVARD dENIVERSITY TROPICAL NEANURIDAE (COLLEMBOLA) Bedos, A. & L. Deharveng, 1991. Cephalachorutes gen. n., a new genus of tropical Neanuridae (Collembola). - Tijdschrift voor Entomologie 134: 145-153, figs. 1-22, tabs. 1-2. [ISSN 0040-7496]. Published 18 December 1991. Cephalachorutes, a new genus of Neanuridae (Pseudachorutinae) Collembola, is described with five new species from Southeast Asia (C. asiaticus sp. n., C. barthae sp. n., C. caecus sp. n., C. centurionis sp. n., C. pestilentiae sp. n.) and one new species from Papua-Niugini (C. murphyi sp. n.). In addition, three species from Africa described in Arlesia, and one species from Nepal described in Pseudachorutella are moved to Cephalachorutes. Correspondence: A. Bedos and L. Deharveng, Laboratoire de Zoologie, Ecologie des Invertébrés terrestres, UPR CNRS 8491, Université Paul Sabatier, 118 route de Nar- bonne, 31062 Toulouse Cedex, France. Key words. - Collembola, Neanuridae, tropical forest. Collections of forest soil fauna from Southeast Asia frequently contain small Pseudachorutinae (Neanuridae, Collembola) of unusual habitus, i. e. with short and wide body and head. Following the classical work of Massoud (1967), these species would have to be placed in either Arlesia or Pseu- dachorutella, according to the number of their eyes. More detailed morphological studies show, how- ever, that, apart from their peculiar habitus, these forms share a set of unique characters which justify to isolate them in a new genus described below. The terminology used is that of Deharveng (1983) for antennae and tibiotarsi; labium setae are named after Massoud (1967); dorsal setae are num- bered after the row-system (Cassagnau 1974); however, the neanurian nomenclature of Dehar- veng (1983) is used when precise setal homologies are uncertain (th. II-III). Abbreviations used in the descriptions: abd., ab- dominal segment; ant., antennal segment; d/m, ratio dens/mucro; S-setae, setae of type ‘S’; th. thoracic segment. The material is deposited in the collection of the Laboratoire de Zoologie, Université Paul Sabatier, Toulouse, France (LEIT) and in the Museum Na- tional d'Histoire Naturelle of Paris, France (MNHN). Cephalachorutes gen. n. Type species: Cephalachorutes asiaticus sp. n. (by present designation). Description of the genus Small size: 0.4 to 0.9 mm. Colour blue to white. Body short and wide (about half as wide as long), juvenile-like (fig. 1); 6th abdominal segment re- duced. Antennae (figs. 2-4). Antennae short and stocky; ant. III and ant. IV fused dorsally. Ant. IV with apical papilla reduced, either indistinct or simple (sometimes feebly trilobed), always fused to the apex of the article; organite ‘or’ globulous, in a ventro-apical position; seven well developed, thickened S-setae on ant. IV (S1 to S4, S7, S8 and MS); S7 much enlarged, overhanging the apex of ant. IV; S4 usually larger than other S-setae but smaller than S7; no s-microchaeta, MS as deve- loped as other S-setae. The ant. III organ has moved to ventral side, with its guard-seta S5 shifted distally towards S8 of ant. IV; S3 close to S2. Ant. I with 7 setae. Head (figs. 7, 9-12). Postantennal organ absent. 8+8 to 0+0 eyes. Buccal cone short and wide. Lines of setae A-B and C-D on labium arranged more or less perpendicularly. Labrum distally enlarged, like planaria-head. Labral chaetotaxy 4/3,4,2 with the 2 setae of the distal row and the lateral setae of the ante-distal row longer than others. Maxilla very thin, needle-like. Mandible thin, with 2 strong ba- sal teeth and a distal comb of 4 to 12 small teeth (the 2 apical ones sometimes slightly larger). Tergites (fig. 14). Ordinary setae very short. S- setae very long and thin, shorter on abd. IV. S-setae 145 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 À Di a . x & . & © - eg Fai x ( ve % x = S i = À N NZ EN / x { i : x i ee | x > nn i a N 2. Fig. 1. Habitus of Cephalachorutes barthae as an exam- ple of the genus. Length: 0.5 mm. arranged after the standard hypogastrurian pattern (2+ms, 2/1,1,1,1,1 on each half-tergite from th. II to abd. V). Sternites and body appendages (figs. 18, 19, 21). 1-2 + 1-2 setae on abd. II sternite. Tibiotarsal chaetotaxyf9atgl CES RO ZA LIZA 6+11 when the seta M is absent. Ventral tube with 3+3 setae. Furcula well developed, tenaculum with 3+3 teeth, dens usually with 6 setae (only 3 setae in C. microphthalma); mucro about half as long as dens, simple, similar to that of Pseudachorutes parvulus. Derivatio nominis. - This genus is called after its unusually large and wide head (from the Greek noun kephale = head). Gender of the genus is masculine. Discussion The definition of Cephalachorutes relies primar- ily on the use of antennal chaetotaxy, which appears once again as an efficient taxonomic tool at suprage- neric level in Neanuridae (Deharveng 1983). Among Pseudachorutinae, Cephalachorutes gen. n. shares with Arlesia Handschin, 1942 and Pseu- dachorutella Stach, 1949 (sensu Massoud 1967) the following characters: postantennal organ absent, maxilla styliform, and mucrodens well developed. Like the species of Arlesia sensu Massoud, the Cephalachorutes species have their seta MS of ant. IV of large size, whereas it is reduced to a micro- chaeta in most Pseudachorutinae and in the prim- itive family Hypogastruridae. In the type species of the genus Pseudachorutella (Pasigillata), this seta is also a microchaeta. Cephalachorutes is easily differentiated from Arlesia and Pseudachorutella by the characters listed in tab.1. These last genera, formerly differ- entiated by the number of eyes (8+8 in Pseudachorutella, less in Arlesia, after Massoud 1967), cannot be separated any more on this ground as the species Arlesia variabilis Thibaud and Massoud, 1983 has 8+8 to 5+5 eyes. In fact, the complex Arlesia-Pseudachorutella, which includes a number of unrelated lines, has to be completely revised. Table 1. Morphological differences between Cephalachorutes and Arlesia/Pseudachorutella complex. Size Habitus Ant. IV apical papilla Position of ant. IV organite “or” S7 (and often S4) of ant. IV vs other S-setae (figs. 3, 5) S2-S3 of ant. III organ Ant. III: $2 length vs S2-S3 distance (figs. 4, 6) Labrum shape Line of labial setae C-D (figs. 7 & 8) Mandible Cephalachorutes less than 1 mm short and wide fused to the apex ventro-apical thicker ventral S2 > [S2-S3] like planaria-head more or less perpendicular to A-B 2 big basal teeth and a distal comb of 4-12 small teeth * observed in Arlesia albipes and in Pseudachorutella asigillata. 146 Arlesia/Pseudachorutella usually more than 1 mm long and relatively narrow not fused, trilobed dorso-apical not thicker external S2 < [S2-S3]* different parallel to A-B different Some of the synapomorphies which define Ce- phalachorutes are unique among Pseudachoruti- nae. They are the habitus, at least in the most evolved species, the fusion of apical papilla to the apex of ant. IV, the position of the ant. IV organite ‘or’ (which has migrated beyond the apical papilla on the ventral side of the article), the hypertrophy of S7 and S4 on ant. IV and the labrum shape. The arrangement of the distal setae of labium is similar to that of Kenyura Salmon, 1954, indicating possi- ble relationships between the two genera. The ven- tral position of ant. III organ is also observed in other genera, but in a different way. It is likely that other characters will prove to be discriminant, such as the maxilla arms structure (figs. 12, 13), but too few species are known in this respect. Characters used in the descriptions of species In addition to the characters, which are tradi- tionally used for the taxonomy of Pseudachoruti- nae, the following chaetotaxic characters were found to vary between species: — seta c3 on head | - number of dorso-external setae on th. II and th. III - seta a2 on abd. IV - seta al on abd. V: 2+2 (al present) or 1+1 setae (al absent) between - S-setae - seta M on tibiotarsus In the literature, the number of S-setae on ant. IV is stated to vary between species, but this is in- correct: there are always 7 S-setae on this article, but usually the thinnest have been overlooked and seta S5 of ant. III has been counted as an ant. IV S- seta. Some of the chaetotaxic differences illustrated on figs. 15-17 might rather be individual variation than real interspecific differences: arrangement of setae is often variable on head and tergites (particularly on abd. VI), with frequent asymmetries. Counting the number of eyes is an other serious difficulty for species identification in Cephalacho- rutes. Observation is easier if animals are treated by KOH before clearing. Check-list of Cephalachorutes — C. minimus (Massoud, 1963) comb. n. (from Arlesia); Angola. - C. delamarei (Murphy, 1965) comb. n. (from Arlesia), Gambia. 7. C.nakaoi (Yosii, 1966) comb. n. (from Pseuda- chorutella); Nepal. - C. microphthalmus (Barra, 1969) comb. n. (from Arlesia); Gabon. 6. C. asiaticus sp. n. (type species of the genus); Thailand, Indonesia, Philippines. BEDOS & DEHARVENG: Cephalachorutes . C. barthae sp. n.; Thailand. . C. caecus sp. n.; Thailand. . C. centurionis sp. n.; Thailand. . C. murphyi sp. n., Papua-Niugini. . C. pestilentiae sp. n.; Thailand. BA UR NU Arlesia pillati Prabhoo, 1971 from India might belong to Cephalachorutes, but its description is insufficient. Distribution, ecology Cephalachorutes occurs in Africa, tropical Asia and Papua-Niugini, but not in tropical America. It is a rather frequent inhabitant of forest soils from sea level up to 3,500 m. Species with 4-8+4-8 eyes were collected from litter, whereas those with re- duced eyes were mostly found in humus and soil. Key to world species of Cephalachorutes Additional characters for the species not redes- cribed in this paper are given in brackets. 1. Dens with 3 setae ....... C. microphthalmus! [3-5+3-5 eyes, traces of pigment, claw un- toothed, distal comb of mandible with 8-10 small teeth] ENSIWIEN'O:Setaei ee eee sree 2 DASTBICYES! AA enk 6. C. astaticus sp. n. MESSICANI SAINT 3 TREE N 4 = essithan TIVE TT 6 4. Distal comb of mandible with 4 small teeth ee do RE onu C. minimus [pigment present, claw untoothed, description unreliable for chaetotaxy] - Distal comb of mandible with more teeth 5 5. Seta a2 present on abd. IV ....... 7. C. nakaoi - Setaa2absentonabd.IV after original drawings OM rina Soren ee TE C. delamarei [pale pigment, claw usually untoothed, some- times claw I with tooth, distal comb of mandible with 11-12 small teeth] GWATEAeyesu ns Si care ie eee eee IO 7 INOS CY ES scone Pe ee 9 Note !: Here could be placed Arlesia pillaii Prab- hoo, 1971: 4 setae on dens, 6+6 eyes, pigment present, claw with tooth, distal comb of mandible with 5-6 small teeth, S7 and S4 both thickened; however, the following characters given in the orig- inal description are not consistent with the defini- tion of Cephalachorutes: 4+4 setae on ventral tube and only one big basal tooth on mandible. 147 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 S8 an Rn | or SG (4 tant ‘ È fi iS N À Figs. 2-13. Differential characters on head between Cephalachorutes and Arlesia. - 2, Apex of ant.IV in C. barthae (ventral side, right antenna; ap: apical papilla; or: organite; circles: sockets of setae); 3, Ant.III-IV of C. centurionis (dorsal side, right antenna; S-setae and sockets of ordinary setae of ventral side in dots); 4, Ant Il organ in C. barthae (ventral side, right antenna); 5, Ant.III-IV of A. albipes (dorsal side, right antenna); 6, Ant.III organ in A. albipes (ventral side, right antenna); 7, Labium in C. centurionis; 8, Labium in A. albipes; 9, Mandible of C. caecus; 10, Mandibles of C. barthae; 11, Labrum in C. barthae; 12, Structure of mouthparts in C. barthae (c: cardo; f: fulcrum, st: stipa; md: mandible); 13, Structure of mouthparts in A. albipes. Scales: 10 u (figs. 2, 9-11), 25 u (figs. 3-8, 12), 50 u (fig. 13). 148 — 242 setae between S-setae on abd. V ....... 8 8. 3+S dorso-external setae on th. II and th. III ARORA AD Roe Me i. 3. C. barthae sp. n. — 2+S dorso-external setae on th. II and th. III at Mai 4. C. pestilentiae sp. n. ORE bIOtarsuskwithiseta ME ta. RARO SLI N nannten 1. C. centurionis sp. n. Tibiotarsus without seta M ...................... 1. Cephalachorutes centurionis sp. n. (figs. 3, 7, 14) Type material. - Holotype, female, Thailand: Chiang Mai province, Doi Inthanon, 2550 m, soil in primary forest, 1.vi.1989, Deharveng and Bedos leg., sample CM 4 (LEIT). — Paratypes, Thailand: 1 male, 1 female, ibid, sample cM 6; 2 males, 2 females, 2 juv., ibid, humus in primary forest, 9.i.1981, Gouze leg., samples THA 93 and THA 94 (7 in LEIT, 2 in MNHN). Description Length 0.48-0.68 mm. Ratio length/width = 2.3. Colour whitish sometimes with slight traces of grey pigment. Antennae. Ant. IV with apical papilla entire, fused to the apex; S-setae large and thick; S7 ba- nana-like and larger than others, overhanging the apex of the segment; S4 smaller than S7 but larger than other S-setae. Setae S2 and S5 of ant. III subequal to S8 of ant. IV; on ant. III, S2 is 5 times as long as S3. Head. No ocelli visible, but sometimes 1+1 min- ute black ocular spots. Seta c3 absent. Mouthparts typical of the genus; distal comb of mandible with 9-10 small teeth. Tergites. On th. II and th. III, 3+S dorso-external setae. On abd. IV, a2 absent. On abd. V, 2+2 ordi- nary setae between S-setae. Body appendages. Seta M present on tibiotarsus. Claw without tooth. Dens with 6 setae. Mucro less than half as long as dens (d/m = 2.1-2.3). Derivatio nominis. - This species is named after our friend Alain Gouze, firemen officer, who col- lected the first specimens of this species (from the Latin noun in genitive case centurio = command- ant). Discussion. - Cf C. caecus. 2. Cephalachorutes caecus sp. n. (fig. 9) Type material. - Holotype, male juvenile, Thailand: Chiang Mai province, Doi Chiang Dao, 950 m, humus in forest, 4.vii.1985, Deharveng leg., sample Dc 67 (LEIT). — Paratypes, Thailand: 1 juv., ibid, 1000 m, soil in forest, BEDOS & DEHARVENG: Cephalachorutes 4.vii.1985, Deharveng leg., sample Dc 56; 1 female juv., 3 juv., ibid, 500 m, wet litter in forest, 17.xii.1980, Dehar- veng leg., sample THA 4 (other specimens in alcohol); 1 female, Chiang Mai province, Doi Mae Tho, 1300 m, humus in forest, 14.vii.1985, Deharveng and Bedos leg., sample cL 11 (4 in LEIT, 2 in MNHN). Description Length 0.55-0.65 mm. Ratio length/ width = 2.2. Pigment absent. Antennae. Ant. IV with apical papilla slightly knobbed, fused to the apex; S7 large, fairly thicker than S4; other S-setae thinner and rather long. Setae S2 and S5 of ant. III are subequal to S8 of ant. IV; on ant. III, S2 is 4.5 times as long as S3. Head. No eyes, no ocular spots of pigment. Seta c3 absent. Mouthparts typical of the genus; distal comb of mandible with 8-9 small teeth, the 2 apical ones relatively stronger. Tergites. On th. II and th. III, 3+S dorso-external setae. On abd. IV, a2 absent. On abd. V, 2+2 ordi- nary setae between S-setae. Body appendages. Seta M absent on tibiotarsus. Claw without tooth. Dens with 6 setae. Mucro more than half as long as dens (d/m = 1.8-1.9). Derivatio nominis. - This species is named for the absence of eyes (from the Latin adjective caecus = blind). Discussion. - C. centurionis and C. caecus are the only species of the genus devoid of eyes. They have also the most developed S-setae on ant. IV and ant. III. They differ by the size of 2 apical teeth of the distal comb of the mandible (stronger in C. caecus), the tibiotarsal seta M (present in C. centurionis, absent in C. caecus) and the ratio dens/mucro (higher in C. centurionis). 3. Cephalachorutes barthae sp. n. (figs. 1, 2, 4, 10, 11, 12, 19) Type material. - Holotype, male, Thailand: Chiang Mai province, Doi Mae Tho, 1300 m, in litter forest, 14.vii.1985, Deharveng and Bedos leg., sample CL 17 (LEIT). — Paratypes, Thailand: 1 female, same sample; 1 male, 2 juv., ibid, sample cL 18; 1 female, Chiang Mai province, Doi Inthanon, 2500 m, humus in primary forest, 2.1.1981, Deharveng leg., sample THA 72; 1 juv., ibid, 2000 m, humus in primary forest, 2.1.1981, Dehar- veng leg., sample THA 77; 2 juv., ibid, 1700 m, litter in primary forest, 1.vi.1989, Deharveng and Bedos leg., sample CM 34 (5 in LEIT, 2 in MNHN). Description Length 0.55-0.70 mm. Ratio length/width = 2. Colour blue. 149 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 x DI Á Pai \ O ; °e | 3 er » 3 VER PA ) cd _@ à > © 2 — A 5 } VE 9 2 14 À N a CÒ 3 De Re ABEL EN. l % °° } \ Hen 2 a ie) J N /p3 \ Age HOV Re Se ce ka Cont ~\ » Bass ger Un A N OUT EN vorn Sl ell Figs. 14-22. Differential characters between species of Cephalachorutes. — 14, Dorsal chaetotaxy of C. centurionis; 15, Ocular area of C. pestilentiae; 16, Ocular area of C. nakaoï; 17, Ocular area of C. asiaticus; 18, Chaetotaxy of leg II in C. asiaticus (ventral side; dotted circles: sockets of dorsal setae); 19, Ventral chaetotaxy of C. barthae; 20, Abd.IV-VI of C. murphyi; 21, Mucrodens of C. asiaticus; 22, Abd.III-VI of C. assaticus. Scales: 10 u (figs. 18, 21), 25 u (figs. 15- 17), 50 u (figs. 14, 19, 20, 22). 150 BEDOS & DEHARVENG: Cephalachorutes Table 2. Differential characters between C. barthae, C. pestilentiae and C. murphyi. barthae Body length/width 2 Seta c3 on head absent Dorso-external setae on th. II-III 3+S Seta a2 on abd. IV absent Seta al on abd. V present Antennae. Ant. IV with apical papilla entire but knobbed, fused to the apex; S-setae rather thin, S4 and particularly S7 thicker. Seta S5 of ant. III shor- ter than S8 of ant. IV; on ant. III, S2 subequal to S5 and 3 times as long as S3. Head. 4+4 eyes, sometimes not obvious. Seta c3 absent. Mouthparts typical of the genus; distal comb of mandible with 8-10 small teeth. Tergites. On th. II and th. III, 3+S dorso-external setae. On abd. IV, a2 absent. On abd. V, 2+2 ordi- nary setae between S-setae. Body appendages. Seta M absent on tibiotarsus. Claw with a minute inner tooth, sometimes indis- tinct. Dens with 6 setae. Mucro more than half as long as dens (d/m = 1.7-1.9). Derivatio nominis. - This species is named to thank Florence Barth for her drawing of the Ce- phalachorutes habitus (fig. 1). Discussion. — Cf C. murphyi. 4. Cephalachorutes pestilentiae sp. n. (fig. 15) Type material. - Holotype, female, Thailand: Chiang Mai province, north of Ban Muang Ngai, 600 m, litter in forest, 4.vi.1989, Deharveng and Bedos leg., sample CM 59 (LEIT). - Paratypes, Thailand: 7 females, 2 juv., same sample (numerous specimens in alcohol) (7 in LEIT, 2 in MNHN). Description Length 0.70-0.85 mm. Ratio length/width = 2. Colour blue. Antennae. Ant. IV with apical papilla knobbed, fused to the apex; S4 and particularly S7 thicker than other S-setae. Setae S2 and S5 of ant. III subequal to S8 of ant. IV; on ant. III, S2 is more than 3 times as long as S3. Head. 4+4 eyes (2+2 ocular spots well distinct in alcohol). Seta c3 present; more setae on lateral area than in other species. Mouthparts typical of the genus; distal comb of mandible with 8-9 small teeth. Tergites. On th. II and th. III, only 2+S dorso- external setae. On abd. IV, a2 present. On abd. V, pestilentiae murphyi 2 2.3 present absent 2+S 25S present absent present absent 2+2 ordinary setae between S-setae. Body appendages. Seta M absent on tibiotarsus. Claw with a minute inner tooth. Dens with 6 setae. Mucro more than half as long as dens (d/m = 1.7- 1.8). Derivatio nominis. - The name of the species refers to its type locality, a doline with a very high level of carbon dioxide up to 5% (from the Latin noun in genitive case pestilentia = bad air). Discussion. - Cf. C. murphyi. 5. Cephalachorutes murphyi sp. n. (fig. 20) Type material. - Holotype, female, Papua-Nuigini: Wau, Bulldog road, 2600 m, in litter of moss forest, 1.x11.1979, Deharveng leg., sample PNG 266 (LEIT). - Paratypes, Papua-Nuigini: 1 male, 1 female, Wau, Bul- ldog road, 2650 m, Rhododendron litter, 1.xii.1979, De- harveng leg., sample PNG 263 (LEIT). Description Length 0.75-0.90 mm. Ratio length/ width = 2.3. Colour blue. Habitus less thick-set than the other species of the genus. Abd. VI not hidden under abd. V. Antennae. Ant. IV with apical papilla knobbed, fused to the apex; S4 and particularly S7 thicker than other S-setae, which are long and thin. Seta S5 of ant. III half as long as S8 of ant. IV; on ant. III, S2 slightly longer than S5 and less than 3 times as long as S3. Head. 4+4 eyes not well distinct. Seta c3 absent. Mouthparts typical of the genus; distal comb of mandible with 9-10 small teeth. Tergites. Dorsal chaetotaxy very reduced, diffi- cult to observe on our material. On th. II and th. III, only 2+S dorso-external setae. On abd. IV, a2 ab- sent and only 1+ 1 setae in the p-row between S- setae. On abd. V, 1+1 ordinary setae between S- setae. Body appendages. Seta M absent on tibiotarsus. Claw with one obvious inner tooth. Dens with 6 setae. Mucro half as long or slightly longer than dens (d/m = 1.8-2). 151 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Derivatio nominis. - This species is named in honour of Prof. Murphy who noticed in 1965 that species from Africa should be separated from the neotropical group of Arlesza. Discussion. - Three species of Cephalachorutes have 4+4 eyes: C. barthae, C. pestilentiae and C. murphyi. They are also devoid of seta M on tibio- tarsus. They can be differentiated after table 2. In addition, C. microphthalmus has 3 to 5 eyes on each side, but the species is distinguished easily from the latter by its dens with 3 setae instead of 6. 6. Cephalachorutes asiaticus sp. n. igs 17182122) Type material. - Holotype, male, Thailand: Chiang Mai province, Ban On Luoi, 500 m, bamboo litter, 20.vii.1985, Deharveng leg., sample CL 44 (LEIT). - Para- types, Thailand: 2 males, 1 female, 5 juv., same sample (other specimens in alcohol) (6 in LEIT, 2 in MNHN). Other material. - The following material has been considered as C. asiaticus: Indonesia: 1 male, Sulawesi Selatan, Maros district, Kappang, Gua Salukkan Kallang (cave), 200 m, flood debris, 14.vii.1986, sample INDO 142; 2 juv., Sulawesi Selatan, Maros district, Kappang, 300 m, in litter of Lantana bushes, 8.vii.1989, Deharveng and Bedos leg., sample SULS 35 (LEIT). - Philippines: 2 males, 3 females, 2 juv., Mindoro Island, Puerto Galera, 150 m, litter in forest, 29.xii.1979, Deharveng leg., sample PHI 116 (LEIT). Description Length 0.55-0.65 mm. Ratio length/ width = 2.3. Colour blue. Antennae. Ant. IV with apical papilla fused to the apex; S7 thicker than other S-setae; S2 slightly larger than S3; other S-setae fairly smaller. Seta S5 of ant. III shorter than S8 of ant. IV; on ant. III, S2 subequal to S5 and less than 3 times as long as S3. Head. 8+8 eyes. Seta c3 absent. Mouthparts typ- ical of the genus; distal comb of mandible with 9- 10 small teeth. Tergites. On th. II and th. III, 3+S dorso-external setae. On abd. IV, a2 present. On abd. V, 2+2 ordinary setae between S-setae. Body appendages. Seta M present on tibiotarsus. Claw with a minute inner tooth. Dens with 6 setae. Mucro half as long as dens (d/m = 2). Derivatio nominis. - This species is named for its large repartition through the tropics in South east Asia. Discussion. - C. asiaticus is the only species of Cephalachorutes with 8+8 eyes. As other primitive character, this species has, with C. nakaoz, the most complete chaetotaxy of tergites in the genus. An- 152 tennal S-setae are relatively poorly developed. Specimens from Indonesia and Philippines have been referred to C. asiaticus, but slight differences in relative width of the body and relative length of mucro were observed with the Thai material. 7. Cephalachorutes nakaoi (Yosii, 1966) comb. n. (fig. 16) Material (type material not examined). - Nepal: 1 male, 1 female, 6 juv., Marsyandi valley, between Chame and Tarapani, 2450 m, in litter and humus of Quercus glauca and Rhododendron, 4.x.77, Deharveng leg., sam- ple Nep 77B-118 (LEIT). Description Length 0.60-0.70 mm. Ratio length/width = 2.2. Colour blue. Antennae. Apical papilla of ant. IV with 3 lobes more or less distinct, fused to the apex; S4 and particularly S7 thicker than other S-setae, which are relatively slender. Seta S5 of ant. III shorter than S8 of ant. IV; on ant. III, S2 slightly longer than S5 and 3 times as long as S3. Head. 5+5 ocelli, sometimes not well distinct. Seta c3 absent. Mouthparts typical of the genus; distal comb of mandible with 7-12 small teeth. Tergites. On th. II and th. III, 3+S dorso-external setae. On abd. IV, a2 present. On abd. V, 2+2 ordinary setae between S-setae. Body appendages. Seta M present on tibiotarsus. Claw with one inner tooth, minute or sometimes indistinct. Dens with 6 setae. Mucro half as long as dens or slightly smaller (d/m = 2-2.2). Discussion. - The original description of the species by Yosii (1966) differs in the apical papilla of ant. IV being conspicuous and distinctly trilobed, whereas it is variable but fused to apex in our material. Other characters are concordant, but chaetotaxic information is lacking for the type specimens which had lost the S-setae on body and antennae, according to the author. C. nakaoz has 5+5 eyes like the African species (C. minimus, C. delamarei and C. microphthalmus). Available dif- ferential characters are few, because the chaetotaxy of these last species is poorly known. The main differences are given in the key. ACKNOWLEDGEMENTS We should like to thank J. M. Thibaud for the loan of Arlesia slides from the Petites Antilles. REFERENCES Cassagnau, P., 1974. Chétotaxie et phylogénie chez les Collemboles Poduromorphes. - Pedobiologia 14: 300- 312. Deharveng, L., 1983. Morphologie évolutive des Collem- boles Neanurinae en particulier de la lignée néanu- rienne. — Travaux du Laboratoire d’Ecobiologie des Arthropodes Edaphiques, Toulouse 4 (2): 1-63. Massoud, Z., 1967. Monographie des Neanuridae, Col- lemboles Poduromorphes a pièces buccales modifiées. - In: Biologie de l'Amérique Australe, CNRS 3: 7-399. BEDOS & DEHARVENG: Cephalachorutes Yosii, R., 1966. Collembola of Himalaya. - Journal of the College of Arts and Sciences, Chiba University 4 (4): 461-531. Received: 22 March 1991 Accepted: 22 April 1991 153 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 BOOK ANNOUNCEMENTS AND REVIEWS Kime, R. D., 1990. A provisional atlas of European My- riapods. Part 1. - Fauna Europaea Evertebrata, Euro- pean Invertebrate Survey, Volume 1: 1-109. Luxem- bourg. [ISSN 0257-7038, obtainable from Musée national d’Histoire naturelle, Luxembourg. Price Lux. Franc 600]. This is the first volume of Fauna Europaea Ever- tebrata, a newly started serial work issued by the European Invertebrate Survey. The E.I.S. promotes the publication of distribution maps on an interna- tional scale. The present atlas is a good example. It is also the first volume of a series of atlasses on the Myriapoda of which nearly 2000 species occur in Europe. This first volume comprises maps for 50 species, all Diplopods, for which the collection of data had been completed and all data checked. The maps are based on published and unpublished re- cords, all from reliable sources, such as collections in natural history museums, publications by authors of repute and recent specialists’ fieldwork. Doubtful records have not been included. Other maps will be published in due course in batches of 50 or more. The European base map with 50 X 50 km squares is used for the purpose, giving appropriate detail for such a large area. Even on this scale not all records could be correctly situated because the locality data, from the literature or on the labels, were too imprecise, a clear hint to collectors and authors to be more accurate. The atlas is an example of international cooper- ation. No less than 40 cooperators from 18 coun- tries are acknowledged for their help with provid- ing records or collections. Despite this seemingly fair coverage of Europe the author recognizes ob- vious gaps in the distributions which are most likely the result of geographically uneven collecting activities in the past. The maps for this good reason have a provisional character. The usual visualiza- tion of older against newer records was not carried through in this project as the records are equally unevenly spread in time. There never were many Myriapod specialists and their collecting activities in a certain area often have a snapshot character with nothing known before or after. The author expresses his hope that the present maps will stim- ulate collecting in those areas which sofar were neglected or undersampled, or were inventoried long ago. The format of the atlas is squarish, an adaptation to the format of the maps. For each species the Opposite page gives relevant ecological informa- tion on habitat and distribution range. Apart from the authorship of the species there are no literature references. A complete bibliography will be pub- 154 lished later as a separate volume. We may call this first volume a must for all Myriapodologists and also for anybody interested in the zoogeography of Europe. [P. J. van Helsdingen] Cherepanov, A. I., 1990. Cerambycidae of Northern Asia. — E.J. Brill, Leiden &c. Volume 1: Prioninae, Distenii- nae, Lepturinae, Aseminae. 642 pp, 296 figs. [ISBN 90 04 09140 8] Hfl. 225.-; volume 2 (in two parts): Cerambycinae. 292 and 354 pp, 133 and 153 figs. [ISBN 90 04 09404 0] Hfl. 270.-; volume 3 (in two parts): Lamiinae. 300 and 308 pp, 120 and 133 figs.[ISBN 90 04 09140 8] Hfl. 290.-. All cloth with dust jacket. A description of the morphology, geographic distribution, and biology of cerambycid beetles (Co- leoptera: Cerambycidae). Keys to taxa based on different developmental stages are presented here for the first time, host relationships examined, and the life cycle and ecology of each species assessed. Volume 1 includes four subfamilies, comprising 124 species, volume 2 includes 108 species, and volume 3 152 species. The data for these books were obtained during field and laboratory investi- gations over wide expanses of Siberia, the Urals and the Far East. The adults of all species, and larvae and pupae of many species are illustrated. The books are translations of the original Russian ‘Usachi Sever- noi Azii’ published from 1979 onwards. It is most fortunate that the knowledge about these beetles has become available in English. Descriptions and biological notes are detailed and certainly useful for e.g. workers in plant protection. Unfortunately, the quality of reproduction (especially of the illustra- tions) is inadequate. [J. van Tol] Veeresh, G. K., B. Mallik & C. A. Viraktamath (eds.), 1990. Social insects and the environment. Proceedings of the 11th International Congress of IUSsI 1990. - E. J. Brill, Leiden &c. 765 pp. [ISBN 90 04 09316 8]. Cloth in dust jacket. Hfl. 150.-. This volume includes 370 papers presented by leading scientists at the 11th Congress of the Inter- national Union for the Study of Social Insects at Bangalore, from August 5-11, 1990. The papers are classified into 30 sections related to the symposia papers of the Congress. These cover various front- iers of research on social insects such as evolution of sociality, polygyny, social polymorphism, kin- recognition, kin-selection, foraging strategies, re- productive strategies, biogeography and phyloge- netics of bees and ants pollination ecology and management of pestiferous social insects. The texts are unrefereed, unedited abstracts of the papers presented and printed as camera-ready copy. [Mainly from information by the publisher] C. L. BELLAMY National Museum of Natural History, Washington, D. C. A REVIEW OF THE PHILIPPINE SPECIES OF THE COROEBUS GORY & LAPORTE GENUS-GROUP (COLEOPTERA, BUPRESTIDAE) Bellamy, C. L., 1991. A review of the Philippine species of the Coroebus Gory & Laporte genus-group (Coleoptera, Buprestidae). — Tijdschrift voor Entomologie 134: 155-176, figs. 1-67. [ISSN 0040-7496]. Published 18 December 1991. The Philippine genera and species of the Coroebus generic group are reviewed. Five species of Coroebus are identified: coelestis Saunders, spinosus (Fabricius), hastanus Gory & Laporte, albiventris sp. n. and bivestitus sp. n. Five species of Cisseicoraebus are recognized: grandis (Kerremans), pullatus (Saunders) comb. n., piperi (Fisher) comb. n., cisseoides (Saunders) comb. n. and bicoloratus sp. n. Philocoroebus gen. nov, is described for azureipennis (Obenberger) comb. n., banahaoensis (Obenberger) comb. n., meliboei- formis (Saunders) comb. n., cyaneoviridis Fisher comb. n., and seven new species: ma- quilingensis sp. n., purpureus sp. n., samarensis sp. n., elongatus sp. n., alius sp. n., adamantinus sp. n. and pseudocisseis sp. n. The first Philippine species of Cissess: aquilonia sp. n. and Hypocisseis: philippinensis sp. n., are described. The second Philip- pine species of Vanroonia: luzonica sp. n. is described. Keys are presented to separate the Philippine genera of the Coroebus group and the species of Coroebus, Cisseicoraebus and Philocoroebus. Correspondence: Dr. C. L. Bellamy, 1651 S. Juniper Street 215, Escondido CA 92025, DAS YA Key words. - Philippines, Buprestidae, Coroebus, taxonomy. The eventual understanding of the genera and relationships within the large, virtually cosmopol- itan tribe Coroebini is dependent upon the correct definition and species placement within or outside the type-genus Coroebus Gory & Laporte. The co- roebine element of the Philippine Islands is partic- ularly exciting due to its position biogeographically as the transition zone between the disparate faunas of southeast Asian, Malesian and Australasian re- gions. Some of the taxa described herein were separ- ated and apparently recognized as new by W. S. Fisher (Department of Entomology, USNM), but he never found the opportunity to realize his apparent intentions following the three main contributions to the Philippine buprestid fauna (Fisher 1921, 192251926): Label data are presented verbatim with commas inserted to separate data from individual lines and the ”/” used to separate data from individual labels. Parenthetical information is added for clarity. The following acronyms are used to indicate the various collections which house and/or loaned material for this study: BMNH, The Natural History Museum, London, England; BPBM, B. P. Bishop Museum, Honolulu, Hawaii, U.S.A.; NMPC, Na- tional Museum, Prague, Czechoslovakia; NSMT, National Science Museum, Tokyo; USNM, National Museum of Natural History, Washington, D.C. and VCLP, Visayas State College of Agriculture, Leyte, Philippines. A few morphometric abbreviations are used in the text as follows: L/W = total length versus width ratio, taken from greatest anterior extension of head to elytral apex; PN W/L = pronotal width versus length ratio; and PS L/W = prosternal length versus width ratio, both using maximum widths and lengths. I was unable to make contact with or secure the loan of material from most of the Philippine col- lections and imagine that there is probably addi- tional material awaiting study and description in the future. It will eventually make sense and be necessary to comment on the phylogenetic relationships of the Philippine Coroebini. However, as there remain numerous undescribed coroebine taxa in the USNM 155 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 1-16. Key figures for Philippine Coroebus generic group, head (frontal aspect), thoracic sternites (ventral aspect), antennae, metatibiae and hing wings. — 1-4, 13, Coroebus hastanus; 5-8, 14, Cisseicoraebus grandis, 9-12, 15, Philocoroebus azureipennis; 16, P. alius sp. n. (scale lines = 1 mm). 156 collections and probably others, e. g. several new species of Brachycoraebus Kerremans from Basilan and Mindanao and new species of both Cryptodac- tylus Deyrolle and Neotoxoscelus Fisher, it is cer- tainly beyond both the scope and intent of this paper. This work has the sole purpose of defining and describing the taxa belonging to the Coroebus genus-group and as a starting point, I have emended the generic key recently presented in the first part of this series (Bellamy 1990). KEY TO THE PHILIPPINE GENERA OF THE COROEBUS GROUP 1. Body generally elongate; pronotum without prelateral carinae; prosternum generally flat- tened, broadly excavated dorsad of procoxae - Body more or less ovoid or elongate ovoid; pronotum with (e.g. figs. 38, 39) or without prelateral carinae; prosternum generally more longitudinally convex and not so excavated 6 2. Mentonniere absent, anterior margin of pros- ternum entire or feebly concave ............. 3 - Mentonniere present, anterior margin of pros- ternum with either a single broad lobe or bi- lobed eA RARA TE 2 eR A CIT RE 4 3. Prosternum with disc produced ventrally, more or less forming a ‘V’ shape, with anterior margin concave between two projected lateral angles; pygidium without median spine and no@wvisıblespas@elytralfapıces mem ee. VREE Fanale 00 Coraebosoma Obenberger — Prosternum more or less entire, anterior mar- gin evenly transverse; pygidium with single projecting medial tooth or spine visible be- tween and past elytral apices (e.g. fig. 25) tane AR PURES De Coroebus Gory & Laporte 4. Head and pronotum uneven, tuberculate SOB ERD OREN Vanroonia Obenberger — Head and pronotum even, entire ............ 5 5. Elytra unicolorous; parts of pronotum, elytra and ventral surface clothed with dense spots, patches or fasciae of white or yellowish pubes- en Cisseicoraebus Kerremans — Elytra with patterns of blue spots and fasciae; pubescence, when present, not in dense con- centrations, much less conspicuous REE Obenbergerula Hoscheck 6. Lateral marginal carinae of pronotum entire, not crenulate or otherwise interrupted … … 7 — Lateral marginal carinae of pronotum crenu- late MATIE PIRES Philocoroebus gen. n. 7. Pronotal lateral carinae, when viewed from above, not visible past premarginal carinae; pronotum more or less flattened ............... RE LI wee ARE AIR. Cisseis Gory & Laporte BELLAMY: Philippine Coroebini — Lateral carinae of pronotum fully visible from above; pronotal disc uneven or convex me- allées, ASO ee Rt 8 8. Pronotum more or less evenly convex me- dially; dorsal coloration apparently mimetic, nitid green with elytral bluish purple spots; elytra without any squamiform setal concen- CHATONS AMENER. RUE Anocisseis Bellamy - Pronotum uneven, with elevated tubercles or costae; dorsal coloration apparently cryptic, somber with irregular iridescent reflections; elytra with irregular concentrations of squami- OE TAS coccosondeaboc Hypocisseis Thomson TAXONOMIC PART Genus Coroebus Gory & Laporte Coroebus Gory & Laporte, 1839: 1. - Baer 1886: 126; Saunders 1871: 104; Kerremans 1903: 231; Schultze 1916: 56; Fisher 1921: 405; Obenberger 1935: 817; Bellamy 1985: 425; 1990: 691. - Type-species: Buprestis undatus Fabricius [as designated: Descar- pentries & Villiers 1967]. Remarks. - The Philippine species of Coroebus have been considered differently in the past. I have decided to narrow the definition in concert and comparison with the type-species and several close relatives from the Palaearctic fauna. This then serves as the starting point and foundation for the subsequent descriptions and new combinations below. Key to the Philippine species of Coroebus 1. Dorsal surface unicolorous, appearing gla- brous, setae very short and sparse, not concen- trated into patches or fasciae (Mindanao) ..... Jer nn C. coelestis Saunders - Dorsal surface with setae concentrated into transverse fasciae or patches ................- 2. 2. Dorsal surface multicolorous, at least when viewed from different angles, some colours, and at some angles, very bright .............. 3 - Dorsal surface unicolorous, ground colour QUitetsOMDELE ee 5 3. Pronotum, more or less, unicolorous, green or deep blue; elytral disc bright green or bluish green, marginal areas black with strong deep blue or violet reflections; elytral with two transverse setose fasciae, one near apical 1/3 and the other just before apical margin (Leyte, Mindanao) ieee OI C. spinosus (F.) — Pronotum, more or less, bicolorous, disc deep green and basolateral portion reddish cupre- ous, cupreous or bright green; elytral disc nitid black, marginal areas in basal 1/2 and entire apical 1/4 green; elytra with a two pairs of setal 157 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 patches on disc of middle 1/3 before single setose fascia at apical 1/3 (Luzon, Mindanao, Negros; Polilloy Samar) ur ee NES SG aida C. hastanus Deyrolle 4. Pronotum subglabrous, disc flattened; dorsal colour blue; elytra with two pairs of white setose patches and two obliquely transverse fasciae (Negros) ......... C. albivestitus sp. n. - Pronotum with long, recumbent setae, disc convex; dorsal colour subnitid black; elytra with white setose patches and one wide fascia past apical 1/3 and a longitudinal band of tes- taceous setae extending along suture (Negros) N N IE C. bivestitus sp. n. Coroebus hastanus Gory & Laporte (figs. 1-4, 13, 18-19) Coroebus hastanus Gory & Laporte, 1839: 10. — Baer 1886: 126; Kerremans 1903: 233; Schultze 1916: 56; Obenberger 1935: 830. Material. - 24 examples (USNM): Luzon, Tayabas Pr., Mt. Dimatabing, IV-V.1925, McGregor, Casiguran; Los Bafios, P.I., Baker; Negros, IV-1911, C.V. Piper; Cuernos Mts., Negros, Baker; Island Polillo, Baker; Island Samar, Baker; Iligan and Davao, Mindanao, Baker. Remarks. - This apparently common and wide- spread species is listed in Obenberger (1935) from Himalaya, Bengale and Japan as well as the Philip- pines. In the USNM collections, there are specimens from Laos, China and Taiwan as well. It may be distinguished from the other Philippine species with its distinct elytral colour pattern shown in fig. 18 and as indicated in the key. The male genitalia are illustrated in fig. 19. Coroebus coelestis Saunders (fig. 17) Coroebus coelestis Saunders, 1874: 319. - Baer 1886: 126; Kerremans 1903: 233; Schultze 1916: 56; Obenberger 1935: 821. Material. - Holotype, female (BMNH): Mindanao. Remarks. - This large, subglabrous species may eventually need to be placed in a distinct genus, but this will await more detailed study of the entire complement of Coroebus. It is quite distinct from the other four species listed here as Philippine congeners and may be separated as in the key. The dorsal habitus is illustrated in fig. 17. Coroebus spinosus (Fabricius) (figs. 20-21) Buprestis spinosus Fabricius, 1801: 214. Coroebus spinosus; Gory & Laporte 1839: 10; Baer 1886: 126; Kerremans 1903: 233; Schultze 1916: 56; Fisher 1921: 406; Obenberger 1935: 837. Coroebus laportei Saunders, 1871: 104. - Baer 1886: 126; Obenberger 1935: 837. Coroebus transversus Kerremans, 1900: 79. - Kerremans 1903: 232; Schultze 1916: 56; Théry 1927: 257; Oben- berger 1935: 837. Figs. 17-21. Coroebus spp., dorsal habitus and dorsal aspect of male genitalia. - 17, C. coelestis; 18, 19, C. hastanus; 20, 21, C. spinosus (scale lines = 1 mm, equal for figs. 19 and 21). 158 Material. - Of transversus, holotype, female (BMNH): Leyte; of spinosus, 5 examples (USNM): Mindanao, Dap- itan and Davao. Remarks. - The type of spinosus was unavailable for study and that of /aportei was not borrowed. I have accepted the synonymy of previous authors, based on determined material in the USNM collec- tion, comparison to the plates in Gory & Laporte (1839) and study of the type of transversus. This beautiful little species is quite distinct from the remainder of the Philippine species and may be separated by its coloration, vestiture and very dif- ferent male genitalia as shown in figs. 20 and 21 respectively. Coroebus albivestitus sp. n. (figs. 22-24) Type material. - Holotype, male (USNM): Philippines, Negros Or., Sibulan, Balingesay, 1981, C. K. Starr. Description Holotype, male. Size, 14.1 X 4.5 mm; elongate, subcylindrical, flattened above; dorsal integument colour nitid black with blue reflection, ventral co- lour subnitid black; head and pronotum sparsely covered with short, stout, recurved black setae ex- cept for slightly more dense patches of testaceous setae as follows: on head one pair dorsal to anten- nal cavities; on either side of pronotum dorsal to marginal carinae near apicolateral angle; elytra moderately setose with setae concentrated into patches and fasciae as in fig. 6; ventral surface partially covered with moderately dense recumbent testaceous setae, especially on hypomera, lateral thoracic sternites, metacoxal plate and first two abdominal sternites. Head: with frontovertex longitudinally de- pressed between large eyes; inner margins of eyes diverging dorsally; circumocular groove extends along anterior margin from near dorsal apex of eye to before ventral apex; each antennal cavity bor- dered dorsally with one arcuate groove; supra-an- tennal grooves confluent with depressed base of frontoclypeus; frontoclypeus in shape of inverted "Y’, with apical margin angularly emarginate; area beneath each eye depressed for reception of basal antennomeres in repose; gena with acute angular projection; antennae with antennomere 2 longer than 3; 3 shorter than 4; 4-10 serrate, longer than wide; 11 shorter than 10, oblong. Pronotum: slightly greater than 1.5X as wide as long; widest at about middle; anterior margin strongly arcuate medially; posterior margin bisin- uate on either side of truncate median portion BELLAMY: Philippine Coroebini anterior to scutellum; basolateral angles obtuse, rounded; lateral margins carinate, finely crenulate, slightly diverging from base to about middle, then roundly arcuate to anterior margin; disc flattened, laterally convex, narrowly explanate before mar- gins; scutellum large, subcordiform, longer than wide; basal margin truncate, lateral angles rounded, apex elongate, acuminate. Elytra: slightly wider at base than pronotal base, widest at about apical third; humeri small, moder- ately elevated, oblique; lateral margins nearly straight from past base to basal third, then widen- ing slightly to apical 1/3 before narrowing to se- parately obliquely truncate apices, apices dentate as in fig. 6; pygidium slightly visible past elytra, broadly arcuate laterally, single short acuminate tooth medially. Fe Underside: suture between first two abdominal sternites feebly indicated laterally; premarginal groove extends along perimeter of sternite 5; femora subfusiform; tibiae flattened, subarcuate; tarsi short with claws bifid, inner teeth pointing inward. Genitalia: as in fig. 24, mounted on a point beneath specimen. Remarks. - Coroebus albivestitus is named for the white setal vestiture of the elytra. This species is immediately distinguished from the remaining Philippine Coroebus spp. with its distinctive colo- ration, elytral setal pattern and male genitalia as illustrated and utilized in the key. Coroebus bivestitus sp. n. (figs. 25-27) Type material. - Holotype, male (BPBM 14609): P. I, Negros Or. Nr. Mt. Talinas, 1020m, 27.VII.1958 / rain forest / H. E. Milliron collector. Description Holotype male. Size, 8.7 X 3.2 mm; elongate, subcylindrical, flattened above; dorsal integument colour subnitid black, with cupreous reflections lat- eral to pronotal gibbosity and along elytral suture, blue reflections in some lateral portions of elytra without dense setal covering; ventral colour sub- nitid black with slight aeneous reflections; head and pronotum moderately to densely covered with recumbent white and testaceous setae, more dense on ventral half of frontovertex and explanate por- tions of pronotum; elytra moderately setose with white setae concentrated into fasciae as in fig. 9 with testaceous setose longitudinal bands, which slightly expand into largest fascia past apical third; ventral surface generally covered with moderately dense recumbent off-white setae. 159 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ur REN IAU ACATENAEEN PRA AACA CONA VICINO] Ae (tet TAT VANS Ain i ayy) N (I en et LL DATENT Tags tll Meers, toy VU) RATA h (1 " Figs. 22-27. Coroebus spp. - 22-24, Coroebus albivestitus sp. n.; 22, dorsal habitus; 23, left lateral aspect; 24, male genitalia, dorsal aspect; 25-27, Coroebus bivestitus sp. n.; 25, dorsal habitus; 26, left lateral aspect; 27, male genitalia, dorsal aspect (scale lines = 1 mm). 160 Head: with frontovertex longitudinally de- pressed between large eyes; a pair of feebly ele- vated small callous-like spots along eye margins near mid-point; inner margins of eyes subparallel, slightly diverging ventrally; circumocular groove extends along anterior margin from near dorsal apex of eye to before ventral apex; each antennal cavity bordered dorsally with one arcuate groove; frontoclypeus in shape of inverted ‘Y’, with apical margin arcuately emarginate; area beneath each eye depressed for reception of basal antennomeres in repose; gena with acute angular projection; an- tennae with antennomere 2 slightly longer, wider than 3; 3 and 4 subequal; 5-10 serrate, longer than wide; 11 subequal to 10, oblong. Pronotum: 1.7X as wide as long; widest at about middle; anterior margin moderately arcuate; pos- terior margin bisinuate on either side of truncate median portion anterior to scutellum; basolateral angles obtuse; lateral margins carinate, crenulate, broadly arcuate from base to apex; disc convex, lateral and basolateral areas explanate before mar- gins; scutellum subcordiform; basal margin sub- truncate, lateral angles rounded, apex elongate, acuminate. Elytra: slightly wider at base than pronotal base, widest at about apical 1/3; humeri small, moder- ately elevated, oblique; lateral margins entirely finely serrate, nearly straight from past base to basal 1/3, then widening slightly to apical 1/3 before narrowing to separately rounded apices; pygidium not visible past elytra. Underside: suture between first two abdominal sternites feebly indicated laterally; premarginal groove extends along perimeter of sternite 5; femora subfusiform; tibiae flattened, subarcuate; tarsi short with claws appendiculate. Genitalia: as in fig. 27, mounted on point be- neath specimen. Remarks. - Coroebus bivestitus may be separ- ated from its congeners with its bicolorous elytral setal pattern and the male genitalia as illustrated in figs. 25 and 27 respectively. The species is named for the bicolored elytral setae. Incertae sedis Coroebus bajulus Deyrolle Coroebus bajulus Deyrolle, 1864: 122. - Baer 1886: 126; Kerremans 1903: 232; Fisher 1921: 406; Obenberger 1935: 820. Remarks. — Saunders (1874) stated that he thought that this species would prove to be a var- iety of hastanus. Although Fisher (1921) listed this species from Negros and Obenberger (1935) also BELLAMY: Philippine Coroebini listed Luzon, I do not believe that this species really exists within the Philippine fauna. The two spec- imens that Fisher recorded are preserved (USNM) within a larger series from several different Philip- pine localities and labelled in his hand as C. hasta- nus; these locality data are listed otherwise irreg- ularly under that species. C. bajalus was described from Ceram and there are very few known distri- butional tracks corroborated by buprestids between this part of Indonesia and the Philippines. Genus Cisseicoraebus Kerremans Cisseicoraebus Kerremans, 1903: 253; Schultze 1916: 56; Fisher 1921: 350; Obenberger 1935: 814; Bellamy 1985: 425; 1990:691. - Type-species: Coroebus gran- dis Kerremans [here designated]. Remarks. - At the time the genus was described, Kerremans (1903) listed three species, C. grandis (Kerremans) from Samar, C. retrolatus (Deyrolle) from Indonesia and C. subcornutus (Fairmaire) from Gabon, with the latter now placed in the genus Exchroaria Obenberger. As Kerremans was generally not in the habit of designating generic type species, and I can find no indication that such has been done for Cisseicoraebus, I designate gran- dis as the type species for this taxon. Key to the Philippine species of Cisseicoraebus 1. Dorsal surface black with setae white and dis- tributed on elytra in small patchs and broad fasciae (CS ES, AoW OE EEEN 2 - Dorsal surface with pronotum green or cupre- ous and elytra green; setae golden and distrib- uted on elytra in patches and narrow fasciae 2. Lateral areas of pronotum and apical 1/2 of elytra densely covered with elongate yellowish white setae; basal 1/2 of each elytron with six setal patches; elytral apices with more elongate slightly recurved denticles; membraneous lobes of parameres more expanded (fig. 37) (Samar) eae ee eee C. grandis (Kerremans) - Lateral areas of pronotum and elytra with a more sparse covering of elongate white setae; both base of elytra and broad fascia from about apical 1/3 to apical 1/6 setose, elytral disc oth- erwise setose as in fig. 28; elytral apices serrate; membraneous lobes of parameres narrow (fig. DO) UZON) MEE C. pullatus (Saunders) 3. Dorsal surface unicolorous, dark green; each elytron with four round and two elongate setal patches, one elongate patch in basal depression (Lamao, Luzon) ............. C. piperi (Fisher) — Dorsal surface bicolorous, head and pronotum differing in colour from elytra; no setal patch 161 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 li si hal I IN? lu, ali aiff | Way N DADI aN nn In) i)! fly! yu) LA (uti \ UN | N "a Mi ij AR DAN YU ya NUO) ) Ir Van MTA à N Ù RO \ I) int] ln TO iin. |) alt | Cu i i il È | | ff i Figs. 28-37. Cisseicoraebus spp., dorsal habitus, male and female genitalia, dorsal aspect. - 28, 29, C. pullatus; 30, 35, C. piperi; 31, C. cisseoides; 32-34, Cisseicoraebus bicoloratus sp. n.; 33, left lateral aspect; 36, 37, C. grandis (scale lines =] mm). 162 in'basal'elytral'depressiontlt ee 4 4. Head and pronotum with cupreous reflection, especially pronotal base; each elytron with five more or less round setal patches (Mindanao) RB C. cisseoides (Saunders) — Head and pronotum roseocupreous; each ely- tron with two round setal patches on disc and two elongate, partially oblique, fasciae on ap- ical 1/3 (Mt. Maquiling, Luzon) ................ ARE Aakn C. bicoloratus sp. n. Cisseicoraebus grandis (Kerremans) (figs. 5-8, 14, 36-37) Coroebus grandis Kerremans 1900: 77. Cisseicoraebus grandis, Kerremans 1903: 253; Schultze 1916: 56; Fisher 1926: 242; Obenberger 1935: 814. Material. - 2 examples (USNM): Island Samar, Baker. Remarks. - This large, attractive species is most similar to C. retrolatus from Sumatra and Borneo. I have not seen an example of retrolatus and, thus, cannot diagnose the differences of these two taxa. Of the Philippine species, grandis is distinctive in its size and pattern of vestiture and may be separ- ated as indicated in the key and as illustrated. Cisseicoraebus pullatus (Saunders) comb. n. (figs. 28-29) Coroebus pullatus Saunders, 1874: 320. - Baer 1886: 126; Kerremans 1903: 223; Schultze 1916: 56; Obenberger 1935: 832. Material. - Holotype, male (BMNH): Luzon. Remarks. - This species is similar in habitus to C. opaculus Obenberger described from India, but I have not been able to compare them directly, so cannot comment on specific differences. At the time this species was described, Saunders (1874) stated that it was ‘most like retrolatus, H. Deyr., of any species I know, which it should follow’. Oben- berger (1935) listed retrolatus under Cisseicorae- bus. Within the Philippine fauna, pullatus is imme- diately distinguished by its coloration, vestiture and male genitalia as indicated in the key and shown in figs. 28 and 29. Cisseicoraebus piperi (Fisher) comb. n. (figs. 30, 35) Coroebus piperi Fisher, 1921: 406. - Obenberger 1935: 832. Material. - Holotype, female (USNM 51490): Lamao, Luzon, PJ. II-VI, 1911, C. V. Piper, collector. BELLAMY: Philippine Coroebini Remarks. - This and the following two species are apparently closely related by virture of a very similar set of integument coloration and the setal patterns of the pronotum and elytra. With no males known for any of the three species, I have illustrated the ovipositors to distinguish C. piperi (fig. 35) from C. bicoloratus (fig. 34). These three species are separated as indicated in the key and by the specific patterns of the elytral setal patterns shown in figs. 30, 31 and 32 respectively. Cisseicoraebus cisseoides (Saunders) comb. n. (fig. 31) Coroebus cisseoides Saunders, 1874: 319. - Baer 1886: 126; Kerremans 1903: 233; Schultze 1916: 56; Oben- berger 1935: 821. Material. - Holotype, female (BMNH): East Mindanao; 1 ex. (NSMT): Agusan d. N., Mindanao, V.1977. Remarks. - At first glance, C. cisseoides appears to be a transitional form between piperi and bico- loratus, however the absense of both males and specimens other than the types makes it difficult to comment further on the relationship of these three species. It may be separated from its congeners as indicated in the key and as discussed previously. Cisseicoraebus bicoloratus sp. n. (figs. 32-34) Type material. - Holotype, female (USNM): Mt. Makil- ing, Laguna, PI, v.9.31 / F. C. Hadden collector; 1 par- atype, female (NMPC): Mt. Maquiling, elev. 1800 m, 24.v.1949, P. Namocale. Description Holotype, female. Size, 12.2 mm X 5.0 mm; elongate, ovoid, flattened above and below; head and pronotum dark cupreous with reddish tinge; elytra bluish green; ventral surface, legs and anten- nae aeneous; dorsal surface moderately densely co- vered with medium-sized shallow punctures, elytra otherwise subrugose; ventral surface sparsely punctate medially, subimbricate laterally; dorsal surface generally with a single short silver recum- bent seta from each puncture, otherwise with dense concentrations of recumbent yellow setae on fron- tal depression between eyes, flattened lateral areas of pronotum and in patches and fasciae on elytra as in fig. 32; ventral surface very sparsely setose medially, much more densely covered with long recumbent yellow setae on hypomera and lateral thoracic and abdominal sternites. Head: frontovertex with lateral lobes produced between eyes, longitudinally depressed medially; 163 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 eyes large, ovoid, with ventral margin truncate dorsal to genal antennal groove, inner margins subparallel; circumocular groove only along inner margins; one small arcuate supraantennal groove with a slight dorsal gibbosity above each antennal insertion; frontoclypeus compressed between large antennal cavities, distal emargination moderately shallow, subtriangular; gena lobe with acute pro- jection. Antennae with antennomere 2 subequal to 1, widest distally; 3 shorter than 2; 4-10 triangu- larly serrate; 11 oblong, curved. Pronotum: 1.66X as wide as long, widest near middle; anterior margin arcuate; posterior margin biarcuate on either side of subtruncate prescutellar median lobe; posterolateral angles subacute; lateral margins broadly arucate from base to apex, crenu- late; disc strongly convex between strongly explan- ate lateral portions. Scutellum large, broadly cordi- form. Elytra: wider than pronotum posterior to humeri, widest at about apical third; lateral margins biar- cuate before gradually narrowing to subtruncate apex; margin crenulate to about apical third then serrulate to apex; premarginally deeply impressed along basal third; disc transversely convex laterally, flattened medially; one basomedial depression on either elytron; epipleuron broadest basally then gradually narrowing along entire length. Underside: prosternum ventrally produced, an- terior margin subtruncate medially, arcuate later- ally with emargination for antennae in repose; process slightly swollen posterior to procoxae then attenuate to rounded apex; metepisternum with inner margin strongly arcuate; metacoxal plate with posterior margin slightly dilated, evenly transverse; sutures between abdominal sternites transverse medially, arcuate laterally; sternite 1 nearly 2X length of 2; 1 + 2 longer than 3 + 4 + 5; 2 5 with groove between pleurite border and disc; 5 broadly arcuate; legs: femora narrow, sub- fusiform; protibiae flattened, meso- and metatibiae subcylindrical; tibiae armed with two short distal spines; tarsomeres 1 4 subequal, each slightly shor- ter than preceeding and with slightly more ex- panded ventral pulvillus; 5 narrow, elongate, claws stout, bifid, inner teeth broad, shorter than outer teeth. Ovipositor: as illustrated in fig. 34, moun- ted on point beneath specimen. Variation. Size, 12.1 mm X 4.8 mm. The single female paratype is virtually identical to the holo- type in all aspects of coloration, vestiture and sculp- ture. Remarks. - This species is named for its bicolor- ous dorsal integument. It comes nearest to C. cis- seoides and may be distinguished as indicated un- der that species and in the key above. 164 Genus Philocoroebus gen. nov. Type-species: Coroebus azureipennis Obenberger [new designation]. Short, ovoid, subcylindrical, convex above, flat- tened below. Head produced, bilobed between eyes; eyes large, inner margins diverging dorsally; fron- tal disc entire; supraantennal transverse groove entire; antennal cavities large; frontoclypeus com- pressed between antennae, distal margin emargi- nate; gena lobate. Antennae serrate from antenno- mere 5. Pronotum wider than long; disc convex, somewhat gibbose; lateral portions explanate, margins carinate; with one short prelateral carina on each side. Scutellum large, triangular or cordi- form. Elytra convex, subparallel, attenuate to separately rounded or angulate apices; epipleuron separated from disc by carina, extending to well beyond mid- dle. Thoracic sternites. Prosternum short, longi- tudinally convex, mentonierre entire, short, feebly produced; process with sides subparallel to rounded apex. Mesepisternum, mesepimeron elon- gate, oblique. Metepisternum narrow, elongate. Metepimeron not visible. Metacoxa short, broad, feebly dilated distally. Abdominal sternites. Suture between sternites 1 and 2 feebly indicated, somewhat concave ante- riorly; sutures between sternites 2, 3, 4 and 5 evenly transverse. Legs. Femora narrowly fusiform. Tibiae elon- gate, with pair of distal spines; metatibia with setal comb on external edge. Tarsi short, tarsomeres 1- 4 subequal, each with ventral pulvillus; 5 narrow, elongate, claws appendiculate. Wing as in figs. 15, 16; radial sector vein extend- ing basally about same distance as median vein; radiomedial crossvein extends from posterior an- gle of radial cell; some branches of anal veins poorly indicated. Genitalia: male aedeagus short, parameres par- tially fused basally, some with various apical mem- branous modifications as shown figs. 42, 44, 46, 48, 50, 52 and 54; ovipositor of ‘normal type’. Remarks. - The generic name is a combination of the prefix from Philippines mated with the nominate generic epithet for this large buprestid tribe. The name was chosen to reflect the fact that this group of species is apparently restricted to various islands in the Philippine archipelago. Fisher (1921) recognized two groups of species in his discussion and key for Coraebus [sic] but did nothing further to separate them. The character state of the pronotal prelateral carina is not abso- lute as some of the species do not exhibit the presence of this state. The loss of the prelateral carina is felt to be a character state reversal as the other aspects of the morphology of these congen- ers agree in the perceived characters of importance. Philocoroebus can be separated from the other members of the tribe as indicated in the generic key above and the species are distinguished below. Key to the species of Philocoroebus 1. Pronotum with prelateral carinae, one on either side’ (e.g. figs: 38, 39)! "2 te tene 2 - Pronotum without prelateral carinae (i.e. figs. AS DIM). na io 9 2Bodyimore ovoid,L/ WEEZE FD. re. 3 - Body more elongate, L/W = 2.50 ........... 6 3. Elytra with transverse setal fasciae ............. EME RRsat he P. azureipennis (Obenberger) — Elytra without setae concentrated into trans- verserfasciaenen le AMA IATA eset meee 4 4. Dorsal surface colour black with pronotum re- flecting blue-green, elytra reflecting deep blue; male genitalia with parameres subparallel and with projecting membranous lobes apically (fig. 48) (Samar) ......... P. samarensis sp. n. — Dorsal surface iridescent blue or blue green; parameres strongly diverging apically, some part of each apex membranous, but without projecting lobe (e.g. fig. 42) i. 5 5. Dorsal integument deep blue; male genitalia as into A2 (LUZOM) rc Beerens ee neee CERES P. banahaoensis (Obenberger) - Dorsal integument blue-green to deep blue; male genitalia as in fig. 44 (Luzon) ............ ne (I ARE P. maquilingensis sp. n. 6. Dorsal integument iridescent green with or withoutibluefreflectio nk rn 7 - Dorsal integument iridescent deep blue with either green or purple reflections ........... 8 7. Elytra with transverse setal fasciae; ventral surface black with slight iridescent reflections (Luzon) mine P. elongatus sp. n. — Elytra wit’ but setae concentrated into trans- verse fae ..e; ventral surface iridescent green RAVAN) I P. meliboeiformis (Saunders) 8. Prosternum with process more narrow in pro- portion to entire length (PS L/W 2.50) (Luzon) bebe est eee P. alius sp. n. - Prosternum broader (PS L/W 2.11) (Minda- MAO) AE EN EAO P. adamantinus sp. n. 9. Pronotum with lateral explanate areas covered with recumbent setae; elytra with small setal patches resembling some spp. of Cisseis (Luzon) st Mers P. pseudocisseis sp. n. — Dorsal surface without any concentrations of Setaelas APOVEL o AEN 10 10. Dorsal surface iridescent green with blue re- flections (Luzon) .... P. cyaneoviridis (Fisher) — Dorsal surface colour deep iridescent purple BELLAMY: Philippine Coroebini (leazon) assess). Be P. purpureus sp. n. Philocoroebus azureipennis (Obenberger) comb. n. (figs. 9-12, 15, 38-40) Coroebus azureus Fisher, 1921: 405 [name preoccupied ] Coroebus azureipennis Obenberger, 1935: 820 [new name] Material. - Holotype, female (USNM 51489): Baguio, Benguet, Luzon, Baker; 8 females (3 USNM, 3 BPBM, 2 CLBC) Mt. Makiling, Laguna, Luzon, P. I., various dates IV-VI.1931 / 1932, F C. Hadden; 1 female (USNM), Agr. College, Laguna, P. I., V.19.[19]31 / F. C. Hadden. Description The original description of Fisher (1921) is more than adequate and only the following diag- nostic comments are given to allow a confident definition of this species in comparison to the many new ones described below. Diagnosis. Holotype, female. Size, 7.3 X 2.9 mm; ovoid, transversely convex above; shining blue green iridescence dorsally; black ventrally; surface generally imbricate; dorsal surface sparsely covered with recumbent grey setae, slightly more dense, regular on elytra; elytral fasciae with white setae as in fig. 38; ventral surface more densely covered with adpressed white setae. Head longitudinally depressed from vertex along entire length of frons; supraantennal grooves strongly arcuate, moder- ately wide; antennal cavities separated by width subequal to each repective width; distal margin of frontoclypeus emarginate, straight medially, angu- late laterally. Pronotum nearly 1.8X wide as long; with prelateral carinae. Prosternum 2.0X as long as wide. Ovipositor as illustrated in fig. 40. Variation (n = 10). Size: length, 5.8 7.3; width, 2.4 2.9 mm; the dorsal coloration varies from steel blue with a slight purple reflection to a deep metallic green without any blue tinge visible. Male: unknown. Remarks. - This species may only possibly be confused with P. elongatus sp. n. due to the similar dorsal coloration and vestiture, however the body proportions and more complex development of the elytral setae pattern of elongatus will immediately separate these two taxa. Philocoroebus banahaoensis (Obenberger) comb. n. (figs. 41, 42) Coroebus banahaoensis Obenberger, 1928: 338. - Oben- berger 1935: 820. Material. - Lectotype, male [new designation] (NMPC 23709): Mt. Banahao, Luzon, G. Boettcher leg.; 1 male 165 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 45 Figs. 38-48. Philocoroebus spp., dorsal and lateral habitus, male and female genitalia. - 38-40, P. azuretpennis, 41, 42, P. bahanaoensis; 43, 44, P. maquilingensis sp. n.; 45, 46, P. purpureus; 47, 48, P. samarensis (scale lines = 1 mm). 166 paralectotype (NMPC 23710): same data; 1 male (USNM), Mt. Banahao, PL, Baker. Description The original description of two syntypes by Obenberger (1928) is sufficient to warrant just the following diagnosis. Diagnosis. Lectotype, male. Size, 5.5 X 2.4 mm; ovoid; convex above; flattened below; dorsal sur- face iridescent greenish blue with moderate purple tinge on elytra; entire surface imbricate; dorsal surface sparsely covered with recumbent grey se- tae, slightly more dense, regular on elytra; ventral surface more densely covered with adpressed white setae. Head longitudinally depressed; supraanten- nal grooves strongly arcuate, wide; antennal cavi- ties separated by distance subequal to their individ- ual widths; frontoclypeus arcuately emarginate distally. Pronotum nearly 1.9X as wide as long, with prelateral carinae; Prosternum nearly 2.3X as long as wide. Genitalia: as in fig. 42, mounted on point with specimen. Variation (n = 3). Size: length, 5.5-5.7; width, 24-29 mm; the dorsal colour varies in both the paralectotype and the only other specimen on hand by both having the purple tint lost completely and thus being mostly steel blue on the elytra. Female: unknown. Remarks. - This species is quite similar to P. maquilingensis sp. n. and other than the differen- ces in the male genitalia, I would have probably left them together. They are separated as indicated in the key and mostly on the basis of the shape and proporational differences of the male genitalia. As there is overlap between these two mountains in both types series, there is undoubtedly some type of ecological allopatry involved which can explain this divergence. Philocoroebus maquilingensis sp. n. (figs. 43, 44) Type material. - Holotype, male (USNM): Mt. Makil- ing, Luzon, Baker; 3 male paratypes: 1 (BPBM), same data as holotype except VI.1.[19]32 / F. C. Hadden collector; 1 (USNM), Lamao, Luzon, III-VI.(19)11 P I / CV Piper Collector; 1 (USNM), Mt. Banahao, P.I., Baker. Description Holotype, male. Size: 6.7 X 2.6 mm; elongate ovoid, convex above, flattened below; dorsal sur- face iridescent bluish green, ventral surface black with strong blue green reflection; head black with blue and purple reflections; pronotum and medial parts of ventrites imbricate, punctate; lateral por- tion of abdominal sternite 1 longitudinally rugu- BELLAMY: Philippine Coroebini lose; elytra rugose; dorsal surface generally moder- ately covered with fine, short, recurved testaceous setae; ventral surface with more dense covering of more elongate, adpressed, white setae. Head longi- tudinally grooved from vertex along entire length of frons; clypeus broadly arcuately emarginate; su- praantennal grooves strongly arucate, relatively broad; antennal cavities separated by distance much less than their separate width. Protonum nearly 1.9X wide as long; short, prelateral carinae feebly indicated. Elytra with lateral carina serru- late; epipleuron extends to apicolateral angle. Prosternum slightly more than 2.4X long as wide; with widely separated, feebly produced, bilobed mentonierre. Genitalia: as in fig. 44, mounted on point with specimen. Variation (n = 4). Size: length, 5.8-6.1; width, 2.4 2.8 mm; the colour of one paratype is a much deeper blue with no trace of green. Female: unknown. Remarks. - Philocoroebus maquilingensis comes nearest to P. banahaoensis and is distin- guished from that species by the differences in coloration, vestiture and the male genitalia. These two species are also close to the next new species, P. samarensis. These species can be separated as indicated in the preceding key. Philocoroebus samarensis sp. n. (figs. 47, 48) Type material. - Holotype, male (USNM): Island Samar, Baker. Description Holotype, male. Size: 4.8 X 2.2 mm; elongate ovoid, convex above, flattened below; dorsal and ventral surfaces shining black with faint blue and purple reflections; pronotum and medial parts of ventrites imbricate, punctate; lateral portion of ab- dominal sternite 1 longitudinally rugulose; elytra rugose; dorsal surface generally moderately co- vered with fine, short, recurved testaceous setae; ventral surface with moderate covering of short, fine, adpressed, white setae. Head longitudinally grooved from vertex along entire length of frons; clypeus convexly emarginate; supraantennal grooves arcuate, narrow; antennal cavities separ- ated by a distance equal to their separate width. Pronotum 1.9X as wide as long; with prelateral carinae. Elytra with lateral carinae finely serrate. Prosternum 1.8X as long as wide. Genitalia: as in fig. 48, mounted on point with specimen. Female: unknown. 167 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Remarks. - This species is unique in that it is both the smallest species and specimen of Philoco- roebus, is entirely black and is the only specimen known from the island of Samar, hence the name. The aedeagus is very different from the other two species of the perceived species-group, P. bana- haoensis and P. maquilingensis and will serve to immediately distinguish P. samarensis from its congeners. Philocoroebus meliboeiformis (Saunders) comb. n. (figs. 49, 50) Coroebus meliboeiformis Saunders, 1874: 321. — Baer 1886: 126; Kerremans 1903: 233; Schultze 1916: 56; Fisher 1921: 406; Obenberger 1935: 831. Material. - Holotype, male (BMNH): C. Luzon. Description Holotype, male. Size: 7.3 mm X 3.0 mm; elon- gate ovoid; convex above, flattened below; shining golden green above and below, except for apical half of elytra which is black; surface generally im- bricate, with elytral disc rugose; moderately co- vered with short, recumbent white setae. Prono- tum 1.6X as wide as long; pronotum with prelateral carinae. Prosternum 2.2X as long as wide. Genitalia: as in fig. 50, mounted on card with specimen. Female: unknown. Remarks. - This is the only species of Philoco- roebus which is brightly coloured on the ventral surface. The holotype is somewhat damaged as it was originally prepared with a very oversized pin; it is now mounted on a card along with the genitalia and the right middle leg. Philocoroebus cyaneoviridis (Fisher) comb. n. (figs. 51, 52) Coroebus cyaneoviridis Fisher, 1922: 14. - Obenberger 1935: 822. Material. - Holotype, male (USNM 24670): Baguio, Luzon, Philippine Islands. Description Fisher’s (1922) original description is quite in- formative, so the following diagnosis is provided solely for comparison. Diagnosis. Holotype, male. Size: 6.9 X 2.7 mm; elongate ovoid, convex above, flattened below; dor- sal surface iridescent green with some lateral and apical blue reflections, epipleuron black with pur- 168 plish blue reflections, ventral surface black; pron- otum and medial parts of ventrites imbricate, punc- tate; lateral portion of abdominal sternite 1 longi- tudinally rugulose; elytra rugose; dorsal surface generally moderately covered with fine, short, re- curved testaceous setae; ventral surface with mod- erate covering of short, fine, adpressed, white se- tae. Head longitudinally grooved from vertex along entire length of frons; clypeus convexly emargi- nate; supraantennal grooves arcuate, narrow; an- tennal cavities separated by a distance equal to their separate width. Pronotum nearly 1.9X as wide as long, without prelateral carinae. Elytra with lateral carinae finely serrulate on basal 2/3, serrate to apicolateral angle, then finely dentate; apices se- parately angulately truncate. Prosternum 2.2X as long as wide. Genitalia: as shown in fig. 52, mounted on point with specimen. Female: unknown. Remarks. - This species and the following are very close in many character states and differ from the remaining congeners in that they are two of three which lack the prelateral carinae. The general morphology is very similar with only the colora- tion of the dorsal integument significantly differ- ent. The male genitalia are very similar yet differ in the details of the projecting membranous lobes. These two species may be separated with their differing coloration. Philocoroebus purpureus sp. n. (figs. 45, 46) Type material. - Holotype, male (USNM): Mt. Makil- ing, Laguna, PI, V.6.[19]31 / FE. C. Hadden collector. Description Holotype, male. Size: 6.2 X 2.4 mm; elongate ovoid, convex above, flattened below; dorsal sur- face iridescent purple with some lateral blue reflec- tions, epipleuron and ventral surface black; head black with blue green reflections; pronotum and medial parts of ventrites imbricate, punctate; lat- eral portion of abdominal sternite 1 longitudinally rugulose; elytra rugose; dorsal surface generally moderately covered with fine, short, recurved tes- taceous setae; ventral surface with moderate cover- ing of short, fine, adpressed, white setae. Head longitudinally grooved from vertex along entire length of frons; clypeus convexly emarginate; su- praantennal grooves arcuate, narrow; antennal cavities separated by a distance equal to their separ- ate width. Pronotum slightly more than 1.8X as wide as long, without prelateral carinae. Elytra with lateral carinae denticulate; apices separately angulately truncate. Prosternum 2.0X as long as wide. Genitalia: as in fig. 46, mounted on point beneath specimen. Female: unknown. Remarks. - The specific epithet is obviously coined for the distinctive dorsal coloration. P. pur- pureus was discussed above in comparison to P. cyaneoviridis. Philocoroebus alius sp. n. (figs. 16, 53-54) Type material. - Holotype, male (USNM): Majayjay, Laguna, Mar. 6, (19)28, Luzon, P.I. / Colln R. C. McGre- gor. Description Holotype, male. Size: 7.8 X 3.1 mm; elongate ovoid, convex above, flattened below; dorsal sur- face iridescent blue green with some laterobasal golden reflections on the pronotum, ventral surface black; head black with blue green reflections on vertex and roseocupreus reflections on frons; pro- notum and medial parts of ventrites imbricate, punctate; lateral portion of abdominal sternite 1 longitudinally rugulose; elytra rugose; dorsal sur- face generally moderately covered with fine, short, recurved testaceous setae; ventral surface with moderate covering of short, fine, adpressed, white setae. Head longitudinally grooved from vertex along entire length of frons; clypeus convexly emarginate; supraantennal grooves arcuate, nar- row; antennal cavities separated by a distance slightly greater than their separate width. Prono- tum nearly 1.7X as wide as long; with prelateral carinae. Elytra with marginal carinae finely serru- late to apicolateral angle, then finely dentate along rounded apical margin. Prosternum 2.5X as long as wide. Genitalia: as in fig. 54, mounted on point with specimen. Wing: as in fig. 16. Female: unknown. Remarks. - Philocoroebus alius comes nearest to P. adamantinus n. sp. and may be separated as indicated in the key above. These species differ in the colour of the dorsal integument, vestiture, the amount of projection of the frontovertex between the eyes, shape of frontoclypeal emargination and in the two type localities, each from one of the two largest and widely separated Philippine islands. The specific name comes from the Latin, meaning ‘another’. Philocoroebus adamantinus sp. n. (fig. 55) BELLAMY: Philippine Coroebini Type material. - Holotype, female (USNM): Butuan, Mindanao, Baker. Description Holotype, female. Size: 6.8 X 2.7 mm; elongate ovoid, convex above, flattened below; dorsal sur- face nitid black with blue reflections on pronotum and elytra and purple reflections on humeri, epi- pleuron and ventral surface black; head black with blue and golden green reflections; pronotum and medial parts of ventrites imbricate, punctate; lat- eral portion of abdominal sternite 1 longitudinally rugulose; elytra rugose; dorsal surface generally moderately covered with fine, short, recurved tes- taceous setae; ventral surface with moderate cover- ing of short, fine, adpressed, white setae. Head with broad longitudinal depression from vertex along entire length of frons; clypeus emarginate; supraantennal grooves evenly transverse medially, arcuate laterally, narrow; antennal cavities separ- ated by a distance slightly greater than their indi- vidual width. Pronotum slightly more than 1.7X as wide as long; with prelateral carinae. Elytra with marginal carinae finely serrulate along basal 2/3, slightly more so on attenuate apical third, but less so on narrow, separately rounded apices. Proster- num 2.3X as long as wide. Male: unknown. Remarks. - The specific epithet is from the Latin, meaning "steel blue”. This species comes nearest P. alius as discussed above. Philocoroebus elongatus sp. n. (figs. 58, 59) Type material. - Holotype, female (USNM): Majayjay, Mar. 30, [19]29, Lagun, Luzon, P. I. / Colln R C McGre- gor; 3 female paratypes (BPBM): 1, P.I. Luzon, Camarines Sur, Mt. Isarug, Pili, 800m, 27.IV.1965 / H. M. Torrevillas collector; 1, Ifugao Prov., Liwo, 8 km E Jayoyao, 1000- 1300m, 30.V.1967 / L. M. Torrevillas collector; 1, same except Jacmal Bunhian, 24 km E Mayoyao, 800-1000m, 27-29.IV.1967 / H. M. Torrevillas, light trap. Other material. - 3 specimens (NSMT): C. Luzon, Mountain Prov., 3. VII.1989. Description Holotype, female. Size: 7.7 X 3.0 mm; elongate ovoid, convex above, flattened below; dorsal sur- face iridescent blue green, ventral surface black; pronotum and medial parts of ventrites imbricate, punctate; lateral portion of abdominal sternite 1 longitudinally rugulose; elytra rugose; dorsal sur- face generally moderately covered with elongate, recumbent grey setae; elytra with elongate, recum- bent white setae concentrated into patches and 169 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 fasciae as in fig. 58; ventral surface with moderate covering of short, fine, adpressed, white setae. Head bilobed, strongly produced anteriorly be- tween eyes, longitudinally grooved from vertex along entire length of frons; clypeus convexly emarginate; supraantennal grooves strongly arcu- ate, wide; antennal cavities separated by a distance subequal to their separate width. Pronotum nearly 1.8X as wide as long; with prelateral carinae. Elytra with marginal carinae finely serrulate to apicolat- eral angle, then finely dentate along rounded apical margin. Prosternum 2.3X as long as wide. Varia- tion (n = 4). In size, 7.7-8.3 X 2.9-3.4 mm; the three paratypes are more strongly green than blue green in the dorsal coloration. Male: unknown. Remarks. - Philocoroebus elongatus is named for being the most elongate member of the new genus. Due to the pubescent patches and fasciae, it may be confused with both P. azureipennis and P. pseudocisseis n. sp. but may be separated from either of these as indicated in the key. P. azureipen- nis is much shorter in proportion and differs in the distribution of the elytral setal patches, while P. pseudocisseis lacks the prelateral pronotal carinae 99 Figs. 49-55, Philocoroebus spp., dorsal habitus and male genitalia. - 49, 50, P. meliboeiformis; 51,52, P. cyaneoviridis; 53, 54, P. alius sp. n.; 55, P. adamantinus sp. n. (scale lines = 1 mm). 170 and has the elytral setae concentrated only in patches, lacking the apical fascia. The three specimens from Mountain Prov., C. Luzon are an apparent variant of this species. They differ by being consistently larger and the colour is more of a brassy green. Without males present, I prefer to neither make these three paratypes nor to consider them as another new taxon. Philocoroebus pseudocisseis sp. n. (figs. 56, 57) Type material. - Holotype, female (USNM): Mt. Makil- ing, Laguna, P. I. IV.21.[19]31 / F. C. Hadden collector; 3 female paratypes: 1 (USNM), same data as holotype; 1 (USNM), same data except V.6.[19]31; 1 (BPBM): same data except IV.19.[19]31 / elevation 3000 ft. / flowers of malaklak. Description Holotype, female. Size: 6.9 X 2.8 mm; elongate ovoid, convex above, flattened below; dorsal sur- face iridescent golden green, epipleuron and ven- tral surface black; pronotum and medial parts of ventrites imbricate, punctate; lateral portion of ab- dominal sternite 1 longitudinally rugulose; elytra rugose; dorsal surface generally moderately co- vered with fine, short, recurved testaceous setae; elytra with elongate, recumbent, stout, white setae concentrated into small patches distributed on disc as in fig. 56; ventral surface with moderate cover- ing of short, fine, adpressed, white setae. Head longitudinally grooved from vertex along entire length of frons; clypeus convexly emarginate; su- praantennal grooves strongly separately arcuate, wide; antennal cavities separated by a distance slightly greater than their individual width. Prono- tum 1.8X as wide as long; lateral area explanate with moderately dense covering of transversely recumbent yellowish white setae; without prelat- eral carinae. Elytra with marginal carinae finely serrulate on attenuate apical third, more finely dentate on apicolateral angle, then entire along separately subtruncate apices. Prosternum slightly more than 2.2X as long as wide. Genitalia: (not illustrated) mounted on a point beneath specimen. Variation (n= 4). In size, 6.9 7.9 X 2.8 3.0 mm; the coloration is fairly constant except for some blue green elytral reflections on the largest paratype. Male: unknown. Remarks. - This species is named for its similar appearance to numerous species of the Australa- sian genus Cisseis but can be separated from that taxon as in the generic key. As discussed above under P. elongatus, P. pseudocisseis comes near both that species and P. azureipennis is general BELLAMY: Philippine Coroebini appearance but differs considerably. These three species may be separated as discussed previously and as in the species key above. Genus Cisseis Gory & Laporte Cisseis Gory & Laporte, 1839: 1. - Kerremans 1893: 117; 1903: 227; Obenberger 1935: 842; Bellamy 1985: 425. — Type-species: apparently not yet designated (see discussion below). Remarks. - In his revision of the Australian species of Cisseis, Carter (1923) briefly discussed the original generic description and the six species assigned by Gory & Laporte (1839). There was no designation in that work of a type-species and Iam unaware of any subsequent designation; this should be left to the eventual, and long overdue, new revision of this largest generic component of the Australian Coroebini. Cisseis aquilonia sp. n. (figs. 60-62) Type material. - Holotype, male (BPBM 14607): Philip- pine Is., Sur, Iriga Camarines, VII.2-14-1932, M. Caneda. Description Holotype, male. Size, 5.5 mm X 2.2 mm; elon- gate, ovoid, transversely subconvex above and below; both surfaces reddish cupreous, head with a golden reflection; head and underside moderately shallowly punctate; pronotum imbricate; elytra ru- gose discally, imbricate laterally; surface with very sparse cover of short, white, adpressed setae; elytra with setae slightly more stout and more dense and with subsquamiform setae in one fascia and one patch on each elytron as in fig. 60. Head: slightly produced between eyes on either side of longitudinal depression of frons; eyes large, inner margins feebly diverging dorsally; ventral arch of eye bordered by moderately deep groove; frontoclypeus with vague transversely sinuate su- praantennal groove; disc constricted between large antennal cavities; distal margin with trapezoidal emargination, oblique laterally, truncate medially; gena grooved beneath eye for antenna in repose, projecting lobe acute. Antennae: antennomere 1 gibbose; 2 slightly shorter than 1, narrower; 3 shor- ter than 2, slightly wider distally; 4 9 subrectangu- lary serrate, length subequal to width; 10 and 11 missing. Pronotum: nearly 2X as wide as long, widest in posterior half; anterior margin biarcuate on either side on medial convexity; posterior margin bisin- uate on either side of median truncate lobe, a pre- 171 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 VI 4 MY TN, Figs. 56-59, Philocoroebus spp., dorsal and lateral habi- tus. - 56, 57, P. pseudocisseis sp. n.; 58, 59, P. elongatus sp. n. (scale lines = 1 mm). marginal carina extends between two arcuations medially; posterolateral angles obtuse, rounded; lateral margins only visible at base when viewed directly from above, carinate, subparallel from pos- terior margin to beyond midpoint, then arcuate to anterior margin; disc flattened, slightly depressed in posterolateral thirds; premarginal carinae nearly entire, not reaching either anterior or pos- terior margin. Scutellum: cordiform. Elytra: slightly wider than pronotum, widest at margin opposite humeri; lateral margins subparal- lel from base to about apical third, then gradually attenuate to separately rounded apices; margins appear serrulate due to lateral imbrications; epi- 172 pleuron very short, only feebly separated from disc; disc feebly transversely convex; each elytron de- pressed at base between humerus and scutellum and declivous past humerus posterolaterally; distal portion of pygidium slighlty visible beyond elytral apices. Underside: prosternum with bilobed menton- niere, process with sides subparallel between pro- coxae, apex broadly rounded; metacoxal plate short, margins subparallel, posterior margin sinu- ate, feebly dilated; abdominal sternites with sutures between 2, 3, 4 and 5 transverse medially, arcuate laterally; length of 1 less than 2 + 3; 1 + 2 slightly longer than 3 + 4 + 5; 2-5 with premarginal groove; 5 with marginal band strongly explanate, a moderately dense fringe of long white recumbent setae on disc before premarginal groove; marginal attenuate laterally, then arcuate to slight median emargination. Leg: femora feebly swollen, sides subparallel; protibia feebly arcuate, slightly flat- tened; metatibia nearly straight, subcylindrical; protarsi with tarsomeres 1-4 each progressively shorter, with ventral pulvilli progressively longer, more expanded; metatarsi with antennomere 1 el- ongate slightly longer than 2 + 3; 5 narrow, with stout simple claws. Genitalia: as in fig. 62, mounted on point with specimen. The holotype is damaged with the fol- lowing parts missing: left antennae, last two anten- nomeres; right antennae, last three antennomeres; left fore leg and middle leg missing most of tibiae and complete tarsi; right middle leg missing entire tibia and tarsus; left hind leg missing entire tarsus. Remarks. - The only revision of Cisseis was by Carter (1923) and was only for the Australian spe- cies. To even discuss this new Philippine species in context of the Australian species and Carter’s revi- sion without considering the remaining comple- ment of Cisseis seems rather moot and such discus- sion would obviously be rather misapplied. However, for comparative purposes, the following discussion may aid the reader in his understanding of this new Philippine species and its, at least superficial, relationship to the Australian congen- ers. C. aquilonia belongs to the group Carter catag- orized as ‘Sect. II, Group B’, those with the elytra vaguely impressed with white pubescence, more or less marbled, but not in circular spots. The key Carter provides for this group is partly based on colour without a clear outcome for aquilonia. A reasonable comparative match is with C. roseocu- prea Hope, which compares well in size and colo- ration, but differs by not having the flattened white elytral setae, but has the lateral pronotal carinae visible from above and the general dorsal sculpture is both denser and the elytra more rugose. The few non-Australian Cissezs spp. listed by Obenberger (1935) will need to be validated as to their correct generic placement in contrast to Anocisseis Bel- lamy (1990), now known from the Philippines and Borneo. Genus Anocisseis Bellamy Anocisseis Bellamy, 1990: 693. — Type-species: Anocis- seis samarensis Bellamy [from original designation]. En y Mt i ÿ Mii V4 : a Al na Ual F [7 u ’ MM 4 set WT ty i lt, 7 vil Hy Wy 63 BELLAMY: Philippine Coroebini Remarks. — This genus and the type-species were fully described in the first part of this series (Bellamy 1990). I have included them here only in context to the generic key and refer the reader to the previous paper for more detail. Genus Hypocisseis Thomson Hypocisseis Thomson, 1879: 49. - Kerremans 1893: 118; 1903: 256; Obenberger 1935: 860; Bellamy 1985: 425; Figs. 60-67. Cisseis, Hypocisseis and Vanroonia spp. - 60-62, C. aquilonia sp. n.; 60, dorsal habitus; 61, left lateral aspect; 62, male genitalia, dorsal aspect; 63, 66, H. azriceps; 63, left dorsal habitus; 66, left lateral aspect; 64, 65, H. philippinensis sp. n.; 64, dorsal habitus; 66, left lateral aspect; 67, V. luzonica sp. n., dorsal habitus (scale lines = 1 mm). 175 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 1988: 417. — Type-species: Hypocisseis laticornis Thomson [from original monotypy |. Cisseoides Kerremans, 1893: 118. - Kerremans 1903: 254; Obenberger 1935: 857; Bellamy 1985: 425. - Type-species: Cisseoides murina Kerremans [from original monotypy]. Remarks. - These taxa were discussed recently by Bellamy (1988). Hypocisseis would benefit from a thorough revision and this new species is des- cribed to encourage such a project. Hypocisseis philippinensis sp. n. (figs. 64, 65) Type material. - Holotype, female (BPBM 14608): P. I., Mindanao, Zamboange del Sur, Lemesahan, 600m, 7.IX.1958 / light trap, H. E. Milliron. Description Holotype, female. Size, 6.6 mm X 2.7 mm; elon- gate, ovoid, transversely subconvex above and below; nitid black with cupreous reflections on imbrications of vertex and pronotum, elytra with aeneous reflection; head with most of frontal area bright roseocupreous; head, pronotum and ventral surface imbricate, elytra imbricate rugose; surface generally sparsely setose with setae short, white, recumbant, those on elytra subsquamiform. Head: large eyes slightly produced on either side of frontal depression; inner margins of eyes subpa- rallel; frontoclypeus with a transverse groove dor- sad; disc constricted between large antennal cavi- ties; distal margin with half-trapezoidal emargination; gena with area along ventral margin of eye excavated, ventral to this transversely grooved for antennae in repose and projecting lobe acute; antennae with antennomere 2 shorter, more slender than 1; 3 shorter than 2; 4-6 triangulary serrate; 7-10 rectangularly serrate; 11 oblong, re- curved distally. Pronotum: width 2X length, widest near base; anterior margin broadly arcuate; posterior margin bisinuate on either side of median prescutellar lobe; basolateral angles obtuse; lateral margins broadly arcuate in posterior half then more straight and converging toward anterior margin; single car- ina on either side extending from posterior margin to about anterior third; disc uneven, with slight depressions in pairs which are filled in by concen- trations of setae as in fig. 64; scutellum broadly cordiform, posterior angle acute. Elytra: slightly wider at humeri than pronotum; humeri moderately elevated; lateral margins roundly acute from base to opposite humeri, the subparallel to about apical third before gradually narrowing to separately rounded apices; margins carinate serrate from near midpoint; epipleuron a 174 small, elongate triangle; disc with slight basal de- pression on either side; setal pattern as in fig. 64; pygidium not visible past apices. Underside: prosternum with bilobed menton- niere; process with sides subparallel between pro- coxae, apex triangularly attenuate; metacoxal plate short, moderately dilated; abdominal sternites 1 + 2 slightly longer than 3 + 4 + 5; sutures between 2,3,4 & 5 evenly transverse; sternite 5 with pre- marginal groove concentric to margin, with a sparse row of elongate testaceous setae projecting toward posterior, apex slightly emarginate. Legs: femora feebly fusiform, ventral surface slightly excavated for tibiae in repose; protibiae slightly arcuate at base, feebly flattened, unarmed at apex; meso- and metatibiae straight, subcylindri- cal; tarsomeres 1 4 each progressively slightly longer, with ventral pulvilli more developed dis- tally; 5 elongate, narrow, claws bifid with inner teeth shorter. This specimen is missing the entire left hind leg. Remarks. - The species listed under Cisseoides by Obenberger (1935) are either from Australia or New Guinea. H. philippinensis represents the first species known from north of the Equator, an ob- vious significant range extension for the genus. I have illustrated the new species in comparison to H. auriceps (Deyrolle) (figs. 63, 66) from New Guinea. These two species differ in their propor- tions, coloration, vestiture, shape and configura- tion of the prelateral pronotal carinae. Genus Vanroonia Obenberger Vanroonia Obenberger, 1923: 29. - Obenberger 1935: 814; 1958: 503; Bellamy 1985: 425; 1988: 416. - Type- species: Vanroonia coraeboides Obenberger [from original monotypy]. Remarks. - This genus and its identity in com- parison to Amorphosoma Laporte was discussed by Bellamy (1988, 1990). The relictual nature of the species of Vanroonia is indicated by the spread of their distribution (i.e. Africa, India, S.E. Asia, Phi- lippines). An eventual revision of Vanroonia would be helpful as an adjunct to that of Oben- berger (1958). Two species are now known from the Philippines as discussed below. Vanroonia marmorea (Deyrolle) Amorphosoma marmoreum Deyrolle, 1864: 127. - Oben- berger 1935: 793; Fisher 1921: 408; 1926: 241. Vanroonia marmorea; Bellamy 1990: 692. Remarks. - This species, as discussed by Fisher (1921, 1926) and Bellamy (1990), is well defined and will be distinguished from its new congener below. Vanroonia luzonica sp. n. (fig. 67) Type material. - Holotype, female (USNM): Quezon Park, Tayabas, P. I, Alt. 1000 ft., VII-1-[19]32 / F.C. Hadden Collector. Description Holotype, female. Size, length 15.0 mm, width (of pronotum) 4.6 mm; elongate, subcylindrical, flattened below; shining black, with some areas of venter, especially epipleuron, reflecting blue pur- ple; head, pronotum and underside generally im- bricate, elytra rugose; surface generally sparsely covered by short recumbent white setae from im- bricate punctures, setae longer and/or more dense in some areas of pronotum and elytra as shown in fig. 67; elytra with short white setae in patterns and stout, semi-erect dark brown setae interspersed on disc (fig. 67, stippled area). Head with frontovertex depressed medially, produced into lateral lobes which project beyond ocular margin; eyes large, subreniform, inner mar- gins feebly sinuate, diverging dorsally; ocular groove extends from slightly beyond dorsal apex along inner margin to before ventral apex, quite wide dorsally; supraantennal groove transverse, bi- arcuate; frontoclypeus compressed between anten- nal cavities, distal margin excavated, straight me- dially, arcuately laterally; gena with large acute projecting tooth ventral to each eye. Antenna with antennomere | stout, geniculate basally; 2 shorter, narrower than 1; 3 shorter than 2; 4-10 serrate, 4 with width subequal to length, 5-10 each with width to length ratio increasing; 11 oblong. Pronotum nearly 2X wider than long, widest at middle; anterior margin arcuate; posterior margin strongly bisinuate on either side of median lobe; basolateral angles obtuse; lateral margins broadly arcuate, crenulate, explanate; disc uneven, subtu- berculate, with biarcuate prelateral carina on either side. Scutellum large, wider than long, anterior margin straight, with angulate lateral lobes, distal lobe acuminate; disc finely rugose. Elytral subequal in width to pronotum, widest at humeri; humeral lobes feebly projecting; one mod- erately deep depression at base between humerus and scutellum on either side; lateral margins car- inate, widely serrate, subparallel from humeri to apical third, then narrowing gradually to broad, separately rounded, dentate apices; disc flattened medially, convexly rounded laterally; epipleuron BELLAMY: Philippine Coroebini widest just beyond base, subparallel to opposite metacoxa, then gradually narrowing to apicolateral elytral angle. Underside. Prosternum shortened, mentonniere angularly bilobed, process slightly compressed be- tween procoxae, attenuately rounded apically; me- tacoxae short, dilated apically; abdominal sternites 1+2 longer than 3+4+5; 5 with preapical groove around entire length, apex straight. Legs. Femora narrowly fusiform; tibiae some- what flattened, with a quadrate cross-section, with two short stout spines on inner margin distally; protibiae arcuate basally; tarsi short, tarsomeres 1 4 subequal, each progressively shorter and with progressively longer ventral pulvillus; 5 narrow, elongate, equal in length to 1 -4 together, claws simple. Genitalia. Ovipositor mounted on card beneath specimen; of ‘coroebine type’. Remarks. - The name is chosen for Luzon, the large northern island of the Philippine archipelago and location of the type locality. This new species can be distinguished from V. marmorea by its larger size, black coloration and elytral vestiture. V. marmorea usually has the two projecting lobes on the frontovertex clothed with dense erect stout testaceous setae. In addition, the head, pronotum and basal two-thirds of the elytra are a dull aeneous colour and most of the thoracic sternites are den- sely covered with adpressed white setae. ACKNOWLEDGMENTS I wish to thank those curators and colleagues who so graciously supplied me with material in their care: S. Bily (NMPC), M. D. Kerley (BMNH), Y. Kurosawa (NSMT) and G. A. Samuelson (BPBM). I also wish to thank Dr. L. Corpuz-Raros (VCLP) for allowing the USNM to retain the holotype of Coro- ebus albivestitus. REFERENCES Baer, G. A., 1886. Catalogue des Coléoptères des Iles Philippines. - Annales de la Société Entomologique de France (6) 6: 97-200. Bellamy, C. L., 1985. A catalogue of the higher taxa of the family Buprestidae (Coleoptera). - Navorsinge van die Nasionale Museum, Bloemfontein 4 (15): 405- 472. Bellamy, C. L., 1988. The classification and phylogeny of the Australian Coroebini Bedel, with a revision of the genera Paracephala, Meliboeithon and Dinocephalia (Coleoptera: Buprestidae: Agrilinae). - Invertebrate Taxonomy 2: 413-453. Bellamy, C. L., 1990. The Philippine Coroebini (Coleop- tera: Buprestidae: Agrilinae) I. Introduction, nomen- clatural changes and descriptions of a new genus and 175 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 species. - Journal of Natural History 24: 689-698. Carter, H. J., 1923. Revision of the genera Ethon, Cisseis and their allies (Buprestidae). - Proceedings of the Linnaean Society of New South Wales 48: 159-176. Descarpentries, A. & A. Villiers, 1967. Catalogue rai- sonne des Buprestidae d’Indochine XIII. Coraebini (3e partie). - Annales de la Société Entomologique de France 3: 471-492. Deyrolle, H., 1864. Description des Buprestides de la Malaisie recueillés par M. Wallace. - Annales de la Société Entomologique de Belgique 8: 1-312. Fabricius, J. C., 1801. Systema Eleutheratorum, vol. 2. - Kiliae, 687 pp. Fisher, W. S., 1921. New Coleoptera from the Philippine Islands. Family Buprestidae, Tribe Agrilini. - Philip- pine Journal of Science 18(4): 349-447. Fisher, W. S., 1922. New buprestid beetles from Borneo and the Philippine Islands. - Proceedings of the Uni- ted States National Museum 61(7): 1-18. Fisher, W. S., 1926. Fauna Samarensis: Coleoptera, Bu- prestidae. - Philippine Journal of Science 31 (2): 235- 244. Gory, H. L. & F L. de Laporte, 1839. Histoire naturelle et iconographie des insectes coléoptères. Monogra- phie des Buprestides. Livraisons 25-36. — Paris [genera paged separately |. Kerremans, C., 1893. Essai de groupement des bupres- tides. - Annales de la Société Entomologique de Bel- gique 37: 94-122. Kerremans, C., 1900. Buprestides Indo-Malais. Troisieme partie. - Mémoires de la Société Entomologique de Belgique 7: 61-93. Kerremans, C., 1903. - In: Wytsman, P. (Ed.) Genera Insectorum, Coleoptera, Fam. Buprestidae, Fasc. 12b; 12c; 12d. pp. 49-338, Verteneuil & Desmet, Bruxelles. 176 Obenberger, J., 1923. Une série de nouveaux genres de Buprestides. - Acta Entomologica Musaei Nationalis Prague 1: 13-44. Obenberger, J., 1928. Opuscula Buprestologica I. Beitrage zur Kenntnis der Buprestiden (Col.). - Archiv für Naturgeschichte 92(A), Heft 9-11 (1926): 1-350. Obenberger, J., 1935. Buprestidae 4. - In: W. Junk, S. Schenkling (Eds). Coleopterorum Catalogus 143:-782- 934. Obenberger, J., 1958. Révision des espèces du genre Van- roonta Obenb. (Co. Bupr.). - Acta Entomologica Mu- saei Nationalis Pragae 32: 503-512. Saunders, E., 1871. Catalogus Buprestidarum Synonymi- cus et Systematicus. - Janson, London, 171 pp. Saunders, E., 1874. Notes on the Buprestidae collected by Professor Semper in the Philippine Islands; with des- criptions of new species. — Transactions of the Ento- mological Society of London 1874: 303-328. Schultze, W., 1916. A catalogue of Philippine Coleoptera. - The Philippine Journal of Science 11: 1-194. Théry, A., 1927. Etude sur les Coléoptères Buprestides appartenant aux collections des grandes musées (1re note: British Museum). - Annales de la Société Ento- mologique de France 96 (3-4): 247-261. Thomson, J., 1879. Typi buprestidarum musaei Thomso- niani, appendix la. - Paris, 87 pp. Received: 25 March 1991 Accepted: 27 May 1991 J. P. DUFFELS Institute of Taxonomic Zoology (Zoological Museum), University of Amsterdam REVISION OF THE GENUS CHAMPAKA (HOMOPTERA, CICADIDAE) FROM BORNEO AND SULAWESI Duffels, J. P., 1991. Revision of the genus Champaka (Homoptera, Cicadidae) from Borneo and Sulawesi. - Tijdschrift voor Entomologie 134: 177-182, figs. 1-7. [ISSN 0040-7496]. Published 18 December 1991. The genus Champaka and its species are redescribed. The genus includes C. celebensis (= C. maculipennis syn. n.) from NW Sulawesi and C. viridimaculata (= C. harveyi syn. n.), a widespread species in Borneo. Lectotypes are designated. The relationships of Cham- paka are discussed. Dr. J. P. Duffels, Institute of Taxonomic Zoology, Department of Entomology, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. During our ongoing studies of the cicadas of Sulawesi, we have noted that the cicada-fauna of this island is highly endemic and a mixture of taxa with Oriental and Australian relationships (Duf- fels 1990). The cicada genus Champaka Distant, 1905, revised in the present paper, has Oriental affinities. The distribution of the genus reflects a biogeographic coherence between Sulawesi and Borneo. The genus Champaka was erected by Distant (1905) for Pomponia viridimaculata Distant, 1889, from North Borneo, which is thus the type-species by monotypy. Later, the same author added two other species to the genus, viz., Champaka harveyi Distant, 1912 from Borneo and Champaka celeben- sis Distant, 1913 from Celebes [= Sulawesi]. Some years later Haupt (1917) described Champaka ma- culipennis from Celebes. The present study reveals the synonymy of the types from Borneo, as well as those from Sulawesi. DEPOSITORIES Abbreviations for depositories of material used in this paper: Natural History Museum (formerly: British Museum (Natural History)), London, Uni- ted Kingdom (BMNH); California Academy of Sciences, Department of Entomology, San Fran- cisco, USA (CAS); Deutsches Entomologisches In- stitut, Eberswalde, Germany (DEI); National Mu- seum of Natural History (formerly: Rijksmuseum van Natuurlijke Historie), Leiden, The Nether- lands (RMNH); private collection Professor Dr. H.J. Müller, Jena, Germany, containing H. Haupt’s Homoptera collection (Coll. Miiller) and Institute of Taxonomic Zoology (Zoological Museum), Uni- versity of Amsterdam, The Netherlands (ZMA). TAXONOMY The references to the genus and the species are a selection of the most relevant literature; further references can be obtained from the catalogues by Metcalf (1962, 1963) and Duffels & Van der Laan (1985). Champaka Distant Champaka Distant, 1905: 60, 70. - Distant 1906: 49, 71; Distant 1912: 39, 56; Moulton 1923: 82, 116, 166. - Type-species by monotypy: Champaka viridimaculata (Distant, 1889). Diagnosis. - Postclypeus globular. Head consid- erably broader than mesonotum. Head about as long as wide between eyes. Male abdomen consid- erably longer than head and thorax together. Lat- eral margins of pronotum ampliated and distinctly toothed at midlength. Male operculum triangular, short, reaching just beyond anterior margin of ab- dominal segment 3 or to half-length this segment. Greatest width of tegmen less than a third of its length. Characterization of the genus Champaka by syn- apomorphies must await phylogenetic analysis of the characters of the species of Champaka and related genera. The combination of the elongate body-shape and the short opercula of the male 10727 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 i Figs. 1-2. Champaka ssp., habitus, males. - 1, C. viridimaculata, Brunei, Badas; 2, C. celebensis, Sulawesi, Palu, 1985. traditionally separates Champaka from other genera. The Champaka species resemble Platylo- mia spinosa (Fabricius, 1787) and related species in the relatively long abdomen, the broad head and the similar body colour and marking, in spite of the long male operculum of these Platylomia species. The structure of the male genitalia is like those found in Platylomia and other related genera. ae to the species of Champaka Tegmen with infuscations at bases of 2nd, 3rd, Sth and 7th apical areas and along posterior longitudinal vein of 7th apical area. Male oper- culum as in fig. 5 and reaching to half length abdominal segment 3 or to almost posterior margin of this segment. Male genitalia as in fig. 3 (Sulawesi) ........ C. celebensis Distant - Tegmen with infuscations at bases of 2nd and 3rd apical areas. Male operculum as in fig. 6 and reaching just beyond posterior margin of abdominal segment 2. Male genitalia as in fig. 4 (Borneo) ....... C. viridimaculata (Distant) Champaka celebensis Distant (es, D5 1) Champaka celebensis Distant, 1913: 80. - Lectotype & 178 [here designated]: ‘N.W.Celebes / (de Giacomi)’ [handwritten], ‘Champaka / celebensis / Dist. type’ [Distant’s writing], ‘Type’ [ print; red margined label], ‘Distant coll. / 1911-383’ [print] (BMNH) [examined]. Champaka maculipennis Haupt, 1917: 305. - Lectotype @ [here designated]: ‘Paloe / N. W. Celebes’, ‘Champaka & / maculipennis Hpt (Collection Miiller) (syn. n.) [examined]. Other material examined: 1 & paralectotype Cham- paka celebensis, N.W. Celebes, de Giacomi, Distant coll. 1911-383 (BMNH); 2 &, Paloe, N. W. Celebes, A. Heyne, Champaka bellicosa Mel. (manuscript name) (DEI). These two specimens may belong to the type series of Champaka maculipennis, which consists of four males from Palu collected by A. Heyne, but the absence of Haupt’s identification labels and the presence of the label with Melichar’s manuscript name refrain me from label- ling these specimens paralectotype; 14, Sulawesi, Palu, 27.x1.1985, at light, J. van Tol (RMNH). Description of the male Ground colour of body brown; ground colour of dorsal surface of head, pronotum and lateral parts of mesonotum olive-green in the specimen col- lected in 1985. Head and thorax silvery pilose with exception of mediodorsal part of pronotum, dorsal surface of pronotal collar and greater part of me- sonotum disk; abdomen dorsally with scattered silvery pilosity in a median and two lateral bands. Underside of body somewhat lighter brown. Head: Postclypeus for the greater part dark cas- taneous with medial, oval, greenish spot at anterior margin and lightbrown lateral parts. A round black spot enclosing the ocelli; three pairs of very small spots are situated laterally of ocelli. Rostrum with dark brown apex passing hind coxae. Thorax: Pronotum with one broad, dark castane- ous, central fascia which widens to the anterior and to the posterior, a pair of juxtaposed or fused, small, black spots in the central fascia just in front of pronotal collar, a pair of fairly large, half-moon shaped, black-brown spots between the two pairs of oblique fissures and two pairs of short, narrow, black-brown lines just above and below proximal ends of posterior oblique fissures. The narrow, black-brown coloration of posterior oblique fis- sures broadens distally and continues along lateral part of ambient fissure as a fascia of variable width. Pronotal collar laterally with dark suffusion. Me- sonotum with vague median fascia, a pair of nar- row, black-brown, slightly converging paramedian fasciae reaching to half the mesonotum length, a pair of small, round, black-brown spots in front of cruciform elevation and a pair of very broad lateral fasciae on posterior half or two-thirds of mesono- tum. Cruciform elevation lighter than mesonotum disk; anterior arms of cruciform elevation with distinct dark mark. Tegmina and wings: Hyaline. Venation ochre- ous to light brown variegated with black-brown; basal cell and extreme basal venation of tegmen of specimen collected in 1985 with light greenish tinge. Tegmina with infuscations around trans- verse veins at bases of 2nd, 3rd, Sth and 7th apical areas; infuscation at base of 7th apical area extend- ing along posterior vein of this area. Lectotype of celebensis with very small, marginal spots at apices of longitudinal veins of tegmina; most of these spots are missing in the other specimens. Legs: Fore femora black-brown with yellowish basal or middle part; middle and hind femora cas- taneous brown, middle femora with black-brown anterior line. Tibiae of all legs and tarsi of fore and middle legs castaneous brown (tarsi of hind legs missing). Operculum (fig. 5): Triangular, light brown, with almost straight lateral and medial margins and a subacutely rounded apex, reaching to half- length abdominal segment 3 or to almost posterior margin of this segment. Abdomen: Castaneous brown dorsally and light brown ventrally, very long, about 1.4 times as long as head and thorax together. Timbal covering with weakly convex anterior margin and weakly con- DUFFELS: Revision of Champaka cave lateral margin Genitalia (fig. 3). Pygofer one and a half times as long as wide; widest at half-length. Both, lateral margins of apical half of pygofer and sides of basal cup of pygofer pronounced. Uncus lobes broad, lateral margins weakly concave, apical margin weakly convex to attenuate laterodistal corner. Measurements in mm (6 @). Body length 49.0- 61.0 (52.9 + 3.8); head width 13.8-15.5 (14.5 + 0.7); pronotum width 14.7-16.4 (15.3 + 0.6); teg- men length 55.7-62.5 (60.5 + 2.3). Distribution (fig. 7) This species was collected in Palu and in an unknown locality in NW Sulawesi. The specimen from Palu in the RMNH was collected from a window-pane of a restaurant in town in 1985. In spite of rather intensive collecting near Palu and in other areas of Sulawesi in recent years, no more specimens of this species became available. Champaka viridimaculata (Distant) (figs. 1, 4, 6. 7) Pomponta viridimaculata Distant, 1889: 421. - Lectotype 6 [here designated] ‘Kina Balu / (Whitehead)’ [handwritten], ‘viridi- / maculata / Dist’ [handwritten], ‘Type’ [print; red margined round label], ‘Distant coll. / 1911-383’ [print] (BMNH) [examined]. Pomponia viridimaculata; Distant 1891: 73, PIX, figs. 9, 9a-b; Distant 1892: xii; Breddin 1900: 180. Champaka viridimaculata; Distant 1905: 66, 70; Distant 1906: 71; Distant 1912: 57, Pl. 6, figs. 49a-c; Distant 1913: 80; Haupt 1917: 306; Moulton 1923: 116, 168. Champaka harveyi Distant, 1912: 57. - Holotype à: ‘Dutch Borneo / Balakpappan / Henry Harvey / 1912-324’ [handwritten], ‘Champaka / harveyi / Dist. type’ [Distant’s writing], ‘Type’ [print; red margined round label] (BMNH) [examined] (syn. n.). Champaka harveyi, Moulton 1923: 116, 168. Champaka viridimaculata harveyt, Haupt 1917: 306. Other material examined. - 1 @ paralectotype Pompo- nia viridimaculata, Kina Balu, Whitehead, Distant coll. 1911-383 (BMNH). Indonesia, Kalimantan: 2 g, Ponti- anak, Borneo Exp., Max Weber (RMNH); 1 6, Sambas, Dr. J. Bosschoo, acq. 1891 (RMNH); 1 8, Sanga Sanga, H. D. Jansen, 1907-203 (BMNH). - Malaysia, Sabah: 2 4, Mt. Kinabalu, Tenom Keningau, 4-8.iii.1964, J. Smart, Royal Soc. Exped., B. M. 1964-250 (BMNH); 1 4, Labuan, 99-279, viridimaculata, Dist. (BMNH); 5 6, Poring, 9 mi. N. of Ranau, 1600’, 26-29.iv.1970, T. W. & M. C. Davies (CAS); 1 Â, Tawau, 10.iv.1970, G. Mendenhall, coll. T. W. Davies (CAS); 3 9, Tawau, Brumas Camp, xi.1974, C. Pruett, B. M. 1975-590 (BMNH); 8 Q, Sandakan Dist., Rumidi, R. Labuk, 16-30.ix.1973, C. Pruett, B. M. 1975- 590 (BMNH). - Malaysia, Sarawak: 1 g, Bidi, 1907-1908, C.J. Brooks (BMNH); 2 6, Kedurong, Moulton, Distant coll. 1911-383 (BMNH); 4 8, Kedurong, iii.1911, J. C. Moulton, 1911-141 (BMNH). - Brunei: 5 4, Brunei, 179 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 area Ar Figs. 3-6. Champaka ssp. — 3, Male pygofer in ventral view, C. celebensis, Sulawesi, Palu, 1985; 4, Idem, C. viridima- culata, Brunei, Badas; 5, Male operculum in ventrolateral view, C. celebensis, Sulawesi, Palu, 1985; 6, Idem, C. viridimaculata, Brunei, Badas. 180 DUFFELS: Revision of Champaka Fig. 7. Distribution of Champaka viridimaculata 110° 115° 120° Waterstradt (BMNH); 2 @, Brunei, booreiland voor de kust (drilling platform off the coast), acq. 1969, C. Kra- nenburg (ZMA); 9 &, Badas, Agathis / swamp forest / secondary vegetation, 27.11.1982, 50-100’, G. S. Robinson, 1982-156 (BMNH); 5 6, Rampayoh R. (north), LP 195, lowland forest, 1-3.iii.1982, 100’, G. S. Robinson, B M 1982-156 (BMNH); 1 4, S. Selanjak, 4464.1432, man- grove, 0 m, 8-9.iii.1984, Maj. T. P. G. Helps, B. M. 1984- 296 (BMNH). Synonymy C. harveyi was separated from C. viridimaculata by the immaculate tegmina, the greyish-white ba- sa! membranes of tegmina and wings and the more pointed male operculum. Study of the holotype of C. harveyi led to the conclusion that this specimen is an immature C. viridimaculata. The holotype of harveyi has very faint, but still recognizable spots on the tegmina. White basal membranes in the tegmina and wings are also found in specimens of C. viridimaculata. The opercula of harveyi seem to be more pointed but this is an artefact due to deformation of the soft body. Description Body brown to dark brown or black-brown with exception of some parts of mesonotum and pro- notum collar, which are light brown or lightly greenish tinged; abdomen of both sexes often cas- taneous. Pilosity on body as in C celebensis but (black dots) and Champaka 125° celebensis (triangle). underside of head, with exception of postclypeus, more thickly covered with long silvery hairs. Head: Postclypeus more globular than in C. cele- bensis; colour dark castaneous with medial, oval, light ochreous to light brownish spot. A black spot encloses the ocelli and a pair of large dark spots is situated between ocelli and eyes. Rostrum dark brown to apex and just reaching posterior margin of hind coxae. Thorax: Pronotum with an, often indistinct, marking consisting of a pair of central fasciae, a pair of spots between the two pairs of oblique fissures and a dark coloration in these oblique fis- sures. Mesonotum shiny and often darker brown than pronotum but without discernible marking. Tegmina and wings: Hyaline. Colour of venation of tegmen as in C. celebensis. Transverse veins of 2nd and 3rd apical areas of tegmen infuscated. A distinct brown spot is found at apex of anterior longitudinal veins of 2nd apical area, while some specimens have two or three more, very small, marginal spots. Basal membranes of tegmina and wings vividly green. Legs as in C. celebensis. Male. Operculum (fig. 6): Triangular, dark brown, with an almost straight medial margin, a weakly convex lateral margin, and an about rectan- gular apex reaching just beyond posterior margin of abdominal segment 2. Abdomen: Castaneous brown to black brown dorsally and somewhat lighter ventrally, very long, 1.3-1.5 times as long as head and thorax together. 181 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Timbal covering distinctly broader than in C. cele- bensis, leaving a smaller part of interior timbal cavity exposed; lateral margin of timbal covering straight or very slightly convex. Genitalia (fig. 4): Pygofer somewhat less than twice as long as wide, widest at half-length. Lateral margins of apical half of pygofer and curved ridges forming sides of basal pygofer cup strongly pro- nounced. Uncus with two lobes narrowing to api- ces. Apex of uncus lobe bidentate, medial teeth of both lobes juxtaposed and slightly recurved, lateral tooth triangular. Female. Operculum: Semicircular shaped and with slightly convex surface covered with scattered pilosity. Operculum reaching just beyond posterior margin of abdominal segment 2. 4 Abdomen: Castaneous brown dorsally and some- what lighter ventrally. Dorsal surface scattered silvery pilose. Abdomen 0.9-1.2 times as long as head and thorax together. Measurements in mm (10 8, 8 9). Body length 8 51.5-56.0 (53.8 È 1.5), 9 36.6-49.2 (42.5 + 4.8); head width & 15.2-15.8 (15.5 + 0.3), 9 12.7-15.8 (14.8 + 1.1); pronotum width & 15.5-17.5 (16.2 + 0.6), Q 12.3-17.1 (15.0 + 1.7); tegmen length 3 51.0-66.2 (63.1 =# 1-7), 9 517-647 (57.6 = 4.6). Distribution (fig. 7) This species is an endemic of Borneo and re- corded from lowland localities only. ACKNOWLEDGEMENTS For the loan of material the author is indebted to the following persons: Dr P. Arnaud and Dr N. D. Penny, San Francisco, USA; Dr H. Gaedike, Eberswalde, Germany; Dr W. Knight, Dr G. S. Robinson and Mr M. D. Webb, London, United Kingdom; Dr J. H. Miiller, Jena, Germany, and Mr J. van Tol, Leiden, The Netherlands. Thanks are extended to Mr D. A. Langerak for preparing the figures, to Mr L. A. van der Laan for making the photographs, and to Mr G. Verlaan for technical assistance. 182 REFERENCES Breddin, G., 1900. Hemiptera gesammelt von Professor Kiikenthal im Malayischen Archipel. - Abhandlungen der Senckenbergischen naturforschenden Gesell- schaft 25: 139-202 + plate 9. Distant, W. L., 1889. Descriptions of new species of Rhynchota collected on or near the Kina Balu Moun- tain, North Borneo. - Annals and Magazine of natural History (6) 3: 419-422. Distant, W. L., 1891. A monograph of Oriental Cicadidae 4: 73-96 + plates vii-ix. - West, Newman & Co., London. Distant, W. L., 1892. A monograph of Oriental Cicadidae 7: i-xiv + 145-158 + plate xv. - West, Newman & Co., London. Distant, W.L., 1905. Rhynchotal notes. xxix. - Annals and Magazine of natural History (7) 15: 58-70. Distant, W. L., 1906. A synonymic catalogue of Homop- tera. Part 1. Cicadidae: 1-207. Distant, W. L., 1912. Homoptera, Fam. Cicadidae, Sub- fam. Cicadinae. — Genera Insectorum 142: 1-64 + plates 1-7. Distant, W. L., 1913. Descriptions of new species belong- ing to the Homopterous family Cicadidae. - Annals and Magazine of natural History (8) 12: 76-81. Duffels, J. P., 1990. Biogeography of Sulawesi cicadas (Homoptera, Cicadoidea). - In: W. J. Knight & J. D. Holloway (eds.), Insects and the rain forests of South East Asia (Wallacea): 63-72. Royal Entomological So- ciety, London. Duffels, J. P. & P. A. van der Laan, 1985. Catalogue of the Cicadoidea (Homoptera. Auchenorhyncha) 1956- 1980. - Series Entomologica 34: i-xiv + 1-414. Haupt, H., 1917. Fiinf neue Homopteren des indo-ma- layischen Faunengebietes. - Stettiner Entomologische Zeitung 78: 303-309. Metcalf, Z.P., 1962. A bibliography of the Cicadoidea (Homoptera, Auchenorhyncha). General catalogue of the Homoptera 8: i-iv + 1-229. - North Carolina State College, Raleigh, N.C. Metcalf, Z.P., 1963. Cicadoidea. General catalogue of the Homoptera 8 (1) Cicadidae: i-vii + 1-585 + 586-919. - North Carolina State College, Raleigh, N.C. Moulton, J.C., 1923. Cicadas of Malaysia. - Journal of the Federated Malay States Museum 11: 69-182 + pls. 1- DI Received: 21 October 1991 Accepted: 24 October 1991 Xe Ze ZHENG Nankai University, Tianjin NEW SPECIES OF THE GENUS MEGACOELUM FIEBER FROM CHINA (MIRIDAE, HETEROPTERA) Li, X. Z. & L. Y. Zheng, 1991. New species of the genus Megacoelum Fieber from China (Miridae, Heteroptera). - Tijdschrift voor Entomologie 134: 183-192, figs. 1-45. [ISSN 0040-7496]. Published 18 December 1991. Seven new species of the genus Megacoelum Fieber from China are described: M. pronotalis, M. chinensis, M. rubripedum, M. pseudopronotalis, M. zout, M. tenuicorne and M. yunnananum. M. fuscescens Hsiao, 1963 is redescribed. Correspondence: Prof. Le Yi Zheng, Dept. of Biology, Nankai University, Tianjin- 300071, P. R. China. Key words. - China; Megacoelum; Miridae; new species. The genus Megacoelum Fieber was established in 1858. Since then more than 60 species have been described, most of them distributed in the Old World tropics and subtropics (Distant 1904, 1910, Oshanin 1910, Poppius 1914, Carvalho 1959, Lin- navuori 1963-1975, Wagner 1970/1971, Kershner 1972). Up to now, only three species were recorded from China. Two of them were described from Taiwan (M. minutum and M. clypeale) by Poppius (1915), and the third (M. fuscescens Hsiao, 1963) was described from Yunnan (Hsiao & Meng 1963). Seven new species, found in China, are described in this paper, and M. fuscescens Hsiao is redescribed. Megacoelum minutum and M. clypeale are only included in the key. A redescription of these species could not be provided since the types were no* accessible to us. In the descriptions, ‘body length’ = the distance from head apex to hemelytral apex, ‘body width’ = the maximum width across both hemelytra at re- pose, ‘head length’ = the length in front view, and ‘pronotum length’ = the length including collar. Measurements are given in millimeters. In the fig- ures of the vesicae, the secondary gonopore is dor- sally directed. The material cited in this paper was all collected in China. The types of the new species are depos- ited at the Biology Department of the Nankai Uni- versity except where mentioned otherwise. TJNM means Tianjin Natural Museum, China. Checklist of Chinese species of Megacoelum Megacoelum pronotalis Li et Zheng sp. n. (Zhejiang, Jiangxi, Shanxi) Megacoelum chinensis Li et Zheng sp. n. (Zhejiang, Fujian) Megacoelum rubripedum Li et Zheng sp. n. (Fujian) Megacoelum pseudopronotalis Li et Zheng sp. n. (Fujian) Megacoelum zoui Li et Zheng sp. n. (Yunnan) Megacoelum tenuicorne Li et Zheng sp. n. (Guangdong, Guangxi, Sichuan) Megacoelum yunnananum Li et Zheng sp. n. (Yunnan) Megacoelum fuscescens Hsiao, 1963. (Guangdong, Guangxi, Hainan, Yunnan) Megacoelum clypeale Poppius, 1915. (Taiwan) Megacoelum minutum Poppius, 1915. (Taiwan) Key to the Chinese species of Megacoelum 1. Cuneus yellow with black apex. Outer basal angle of membrane with a whitish spot. Body small, length 5mm ......... minutum Poppius — Cuneus at least basal 2/3 black, membrane uni- colorouslyidarkismoky) rm RAR 2 2. Black, with calli and areas before and behind reddish brown. Pronotal basal margin nar- rowly yellow, embolium and outer cuneus red- dish, apical third of cuneus yellowish brown MET a RAI: AO yunnananum sp. n. - Calli concolorous with pronotum. Hemelytra differentlygcolourec gn 3 3. Pronotum medially with black markings … 4 183 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 1-5. Megacoelum fuscescens Hsiao. - 1, vesica in dorsal view; 2, spicule; 3, 4, left clasper, 5, right clasper. - Pronotum without black markings .......... 6 4. Pronotum with a triangular black spot at ante- rior margin, a round median black spot at hind margin of calli, and a long black spot at hind margin. Rostrum only reaching to apex of me- SOcOxae me, clypeale Poppius - Pronotum with only one percurring median blackimarkinigg get toe 5 5. Pronotal median black marking broad and well-defined. Pubescence sparse and long PANELEN dux pronotalis sp. n. - Pronotal median black marking vague and slender: Pubescence very dense and short AM E RE NE Ca z0u1 SP. n. 6. Pubescence dense and short. Pygofer without a knob before left clasper ... rubripedum sp. n. — Pubescence sparse and long. Pygofer with a knobibetorenlefadlasper rn ea 7 7. Pronotal lateral margin and basal portion dar- kened, extreme basal margin yellowish brown DAR MN ne N. fuscescens Hsiao =WPronotumlunicoloroussaar re: 8 8. Frons with reddish transverse stripes. Clypeus somewhat prominent. Antennae slender DE SL EE LARA tenuicorne sp. n. - Frons without light-colored stripes ......... 9 9. Knobon pygofer before left clasper very short. Body comparatively short and broad. Length 688587 A. an ne EN chinensis sp. n. - Knob on pygofer before left clasper large and sharp. Body slender, length 8.37-8.56 REI OSO cris cle sean pseudopronotalis sp. n. 184 IN DESCRIPTIONS 1. Megacoelum pronotalis sp. n. (figs. 6-10) Type material. - Holotype male: Mt. Tianmu (30° 24 N, 119°30'E), Zhejiang Prov., 15.viii.1965, L. C. Wang leg. - Paratypes: 5 males, 3 females, with same data as holotype; 12 males, 19 females (TJNM), same data as holotype, S. L. Liu leg.; 6 males, 1 female, Mt. Lushan (29° 24’N, 115° 54’E), Jiangxi Prov., 22/24.vii.1957, S. H. Ying leg.; 1 male, 5 females (TJnM), Ku-ling (29° 24'N, 115°54’E), Jiangxi Prov., 10. vii/7.viii.1935, O. Piel leg.; 2 males, Jiangxi Prov.; 1 male, Zhenba County (32°30’N, 107° 54E), Shanxi Prov., 20.vii.1985, S. Z. Ren leg. Description. - Dark brown, female paler than male, with long and sparse brown hairs. Head dark brown, juga faintly reddish; antennal segment I, IV and extreme base of II dark brown; II brown, grad- ually turned into dark brown or black apically; III and extreme base of IV yellowish brown. Rostrum surpassing the hind coxae slightly. Collar dirty dark brown, slightly thinner than the basal diameter of antennal segment II, with a row of long black hairs. Pronotum shiny brown, with a darker broad medial longitudinal stripe, sometimes completely dark brown or black (especially the males), or com- pletely brown (especially the females), slightly wrinkled transversely, calli smooth. Scutellum dark brown to black, slightly convex, finely wrinkled transversely, with a transverse impression before extreme apex; pubescence sparse, brown. Mem- brane dirty black. Legs dark brown, tibial spines shorter than the tibial diameter. Thoracic ventral surface dull, dirty black, hairless. Venter shiny, un- evely reddish brown, male pygofer with a long knob (length 0.33) before left clasper (fig. 6). Ves- ica with 3 shagreened membranous lobes, an elon- gate sclerotized plate, connected with a flossy membranous lobe topped with dense minute hairs (figs. 9, 10); left clasper much longer than the right one (figs. 6-8). Measurements: Body length 8.56-10.09, width 2.93-3.30; head apex to cuneal suture 6.14-7.70. Head length male 1.32:1.35, female 1.27-1.32; width male 1.35-1.39, female 1.35-1.42. Vertex width male 0.39-0.40, female 0.51-0.52. Antennal segment lengths male 1.32-1.40 : 3.42-3.68 : 2.41- 2.58 : 1.20-1.29, female 1.31-1.42 : 3.65-3.95 : 2.61- 2.81 : 1.47-1.61. Rostrum length 3.75-3.89. Pron- otum length 1.57-1.68, width 2.55-2.91. Scutellum length 1.40-1.47. Corium length 4.36-4.69. Length of anterior femur : tibia : tarsus = 2.34-2.42 : 2.75- 2.93 : 0.74-0.82; length of hind femur : tibia : tarsus = 2.89-4.15 : 6.03-6.70 : 0.70-1.04. Remarks. - M. pronotalis differs from other known species in the large size, characteristic markings on the pronotum, long and sparse hairs, and the structure of genitalia. 2. Megacoelum chinensis sp. n. (figs. 11-17) Type material. - Holotype: Male, Sangang (27° 42’N, 117°36 E), Chong’an County, Fujian Prov., 22.vi.1965, L. C. Wang leg. - Paratypes: 3 males, 3 females, with same locality as holotype, 22/23.vi.1965; 3 males, 8 females, with same locality as holotype, 20.vii./8.viii.1982, H. G. Zou et al leg.; 2 males, Guadun (26°42’N, 117°36'E), Chong’an County, Fujian Prov., 6.viii.1982, C. Chen & P. P. Chen leg.; 1 male (TJNM), Aotou (27° 18'N, 118°6’E), Jianyang County, Fujian Prov., 21.vi.1965, S. L. Liu leg.; 1 male (TJNM), Mt. Tianmu (30° 24’N, 119° 30’), Zhejiang Prov., 15.viii.1965, S. L. Liu leg.; 1 male (TJNM), with same data as holotype, S. L. Liu leg. Description. - Body relatively short and broad, dark brown to black. Head dull, unevenly brownish black. Antennal segment I thickest, brown, ventral surface darker, with several black setae on inner surface; II brown, slightly incrasste and darkened into black apically, extreme base black; III and IV reddish brown, with paler extreme base, as thick as the base of II. Collar dirty black, slightly thinner than the basal diameter of antennal segment II, with a row of elongate and slender pale hairs. Pronotum shiny, black with pale sparse elongate pubescence, wrinkled transversely, calli smooth. Scutellum slightly convex, densely and finely wrinkled transversely. Hemelytra with sparse long Li & ZHENG: New species of Megacoelum pale pubescence, membrane smoky black. Legs brown, apical 1/6 of femora darker, hind tibiae dark brown. Thoracic ventral surface dirty dark brown. Venter reddish brown, male pygofer with a very short knob before left clasper. Vesica with 4 large membranous lobes, two of which with ap- ical fine shagreened teeth (fig. 16), a slender rod- like sclerotized spicule curved before apex (fig. 17), and an angulate lobe densely and strongly sha- greened (fig. 15). Left clasper (figs. 11-13) long and curved, right clasper short and thick (fig. 14). Measurements: Body length 6.88-8.74, width 2.48-3.15; head apex to cuneal suture 5.12-6.75. Head length 1.09-1.16, width 1.20-1.32. Vertex width male 0.46-0.50, female 0.49-0.50. Antennal segment lengths 1.21-1.42 : 3.02-3.35 : 2.28-2.68 : 1.21-1.41. Rostrum length 3.15-3.38. Pronotum length 1.27-1.44, width 2.08-2.48. Scutellum length 1.01-1.17, width 1.07-1.24. Corium length 3.45- 4.22. Length of anterior femur : tibia : tarsus = 1.68-2.09 : 2.41-2.65 : 0.65-0.67, length of hind femur : tibia : tarsus = 3.08-3.80 : 4.86-5.43 : 0.67- 0.78. Remarks. - Body shape and colour somewhat similar to M. pronotalis sp. n., but differs in smaller size, longer pubescence and the different structure of male genitalia. 3. Megacoelum rubripedum sp. n. (figs. 19-25) Type material. - Holotype male: Sangang (27° 42’N, 117°36' E), Chong’an County, Fujian Prov., 5.viii.1982, H. G. Zou leg. - Paratypes: 3 males, 2 females, with same locality as holotype, 29.vii/7.viii.1982, C. Chen et al leg.; 5 males, 2 females, Hexi (24°48’N, 117°12’E), Nanjing County, Fujian Prov., 22.vii.1965, L. C. Wang leg. Description. - Reddish black, sometimes the fe- male slightly paler,with dense and short yellowish brown hairs. Head dark brown,dull, antennae slender and elongate, red to brown, extreme base of segment II black, extreme base of III, extreme base and apex of IV yellowish brown. Collar brown, thinner than the basal diameter of antennal seg- ment II, with a row of elongate hairs with swollen base. Pronotum black, basal margin narrowly brownish red, densely and transversely wrinkled, calli indistinct. Scutellum black, with fine and dense transverse wrinkles. Hemelytra black, embolium and cuneus brown to red; membrane smoky black, apical angle of large cell nearly rectangular. Legs reddish brown to red. Thoracic ventral surface in- cluding coxae dull black. Venter unevenly brownish black, with sparse brown hairs. Male pygofer with- out any knob before left clasper. Vesica (fig. 22) with 4 membranous lobes (one of them strongly 185 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 6-17. Megacoelum spp. - 6,7, Megacoelum pronotalis sp. n., left clasper; 8, idem, right clasper; 9, idem, vesica in dorsal view; 10, idem, vesica, sclerotized plate and flossy membranous lobe, ventral view; — 11-13, Megacoelum chinensis sp. n., left clasper; 14, idem, right clasper; 15, idem, vesica in dorsal view; 16, idem, vesica, shagreened lobe; 17, idem, vesica, spicule. 186 and roughly shagreened apically, as fig. 24) and dorso-medially a mushroom-like appendage with a sclerotized cap-shaped apex (figs. 23, 25). Left clasper elongate and curved (figs. 18-20), right clasper short, broadly toothed medially (fig. 21). Measurements: Body length 5.81-7.63, width 2.05-2.68; head apex to cuneal suture 4.37-6.03. Head length 0.99-1.22, width 1.09-1.32. Vertex width male 0.43-0.45, female 0.41-0.51. Antennal segment lengths 1.11-1.51 : 2.60-3.75 : 2.28-3.02 : 1.17-1.41. Rostrum length 3.28-3.35. Pronotum length 1.10-1.40, width 1.80-2.21. Scutellum length 0.90-1.21, width 0.90-1.21. Corium length 2.93- 3.82. Length of anterior femur : tibia : tarsus = 1.58-1.89 : 1.91-2.48 : 0.54-0.63, length of hind femur : tibia : tarsus = 2.75-3.52 : 4.14-5.26 : 0.59- 0.76. Remarks. - Differs from other known species in the slender and elongate antennae, red or reddish brown legs, dense and short pubescence and the structure of male genitalia. Li & ZHENG: New species of Megacoelum 4. Megacoelum pseudopronotalis sp. n. (figs. 26-30) Type material. - Holotype male: Yanshan (27° 18'N, 117° 18’E), Shaowu County, Fujian Prov., 28.v.1965, L. C. Wang leg. - Paratypes: 1 male, with same data as holo- type; 1 male (TJNM), Sangang (27°42’N, 117°36’E), Chong’an County, Fujian Prov., 22.vi.1965, S. L. Liu leg. Description. — Black or reddish brown, with sparse dark brown hairs. Head dark brown to black, weakly shining. Gula horizontal. Antennal seg- ment I, extreme base of II black; basal half of II yellowish brown, III and IV absent. Collar dull, black, as thick as the basal diameter of antennal segment II, with a row of elongate dark hairs. Pronotum black, shiny, pubescence sparse, weakly wrinkled transversely, calli smooth with margins indistinct. Scutellum shiny black, finely and trans- versely wrinkled. Hemelytra dark reddish brown, membrane smoky black. Legs brownish black, an- Figs. 18-25, Megacoelum rubripedum sp. n. - 18-20, left clasper; 21, right clasper; 22, vesica in dorsal view; 23, vesica, lateral view of mushroom-like lobe; 24, same, top view; 25, vesica, shagreened lobe. 187 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 TRA, Pi; il N MG N Wey “1 Ty EML 4 Ù e (OLI, + Ni ag \ N \ N \ i Figs. 26-35, Megacoelum spp. - 26, 27, Megacoelum pseudopronotalis sp. n., left clasper; 28, idem, right clasper; 29 idem, vesica in dorsal view; 30, idem, vesica, sclerotized plate and flossy membranous lobe; 31-33, Megacoelum zoui sp. n., left clasper; 34, idem, right clasper; 35, idem, vesica, dorsal view 188 terior and middle tibiae excepting extreme apex brown. Thoracic ventral surface dull, black. Venter brownish black. Knob on pygofer before left clasper in male large and sharp, length 0.39 (fig. 26). Vesica (fig. 29) very similar to that of M. pronotalis sp. n., but the sclerotized plate more strongly hooked apically (figs. 29, 30). Apical 1/3 of the left clasper (figs. 26, 27) twisted. Right clasper (fig. 28) close to that of M. pronotalis sp. n. Measurements: Body length 8.37-8.56, width 2.75-2.80; head apex to cuneal suture 5.91-6.14. Head length 1.24-1.25, width 1.34-1.35. Vertex width 0.46-0.48. Lengths of antennal segment I and I = 1.34-1.35 : 3.35-3.75. Rostrum length 3.73- 3.76. Pronotum length 1.38-1.39, width 2.41-2.43. Scutellum length 1.16-1.22, width 1.29-1.32. Co- rium length 4.00-4.09. Length of anterior femur : tibia : tarsus = 2.18-2.21 : 2.61-2.66 : 0.70-0.72, length of hind femur : tibia : tarsus = 2.69-2.75 : 5.70-5.90 : 0.80-0.81. Remarks. - Closely allied to M. pronotalis sp. n. in the body size and shape, but the pronotum al- ways black, the structure of vesica and the left clasper different. 5. Megacoelum zoui sp. n. (figs. 31-35) Type material. - Holotype, male, Mengxiu, Ruili County (24°0'N, 97°48'E), Yunnan Prov., 3.1x.1979, H. G. Zou leg. - Paratypes: 1 male, 1 female, with same locality as holotype, 2.ix.1979, J. X. Cui and G. Q. Liu leg. Description. - Dark brown, weakly shiny with very dense and short brown pubescence. Head blackish brown, clypeus slightly convex; antennal segment I, extreme base of II and the apical parts of III and IV dark brown, darkened apically, bases of III and IV yellowish brown. Collar brown, me- dian part black, thinner than the basal diameter of antennal segment II. Pronotum medially with a vague longitudinal black stripe. Scutellum convex, black or dark brown, extreme apex brown. Mem- brane smoky black, veins black. Front legs brown, with darker femora; middle femora and the basal half of tibia dark brown; hind leg reddish black. Thoracic ventral surface dark brown. Venter dark reddish with brown hairs. Knob on pygofer before left clasper in male small. Vesica (fig. 35) with 4 membranous lobes, one of them strongly and den- sely toothed apically. Left clasper (figs. 31-33) long and curved, right one (fig. 34) short and straight. Measurements: Body length 7.81-8.37, width 2.61-2.79; head apex to cuneal suture 5.57-5.59. Head length 1.09-1.14; width male 1.25-1.27, fe- male 1.25. Vertex width male 0.43-0.46, female Li & ZHENG: New species of Megacoelum 0.46. Antennal segment lengths 1.14-1.31 : 3.08- 3.35 : 2.27-2.51 : 1.17-1.21. Rostrum length 3.35- 3.72. Pronotum length 1.30-1.32, width 2.25-2.31. Scutellum length 1.12-1.22, width 1.13-1.26. Co- rium length 3.58-3.86. Length of anterior femur : tibia : tarsus = 2.01-2.11 : 2.55-2.58 : 0.60-0.67, length of hind femur : tibia : tarsus = 3.15-3.25 : 5.16-5.49 : 0.70-0.80. Remarks. - This species is distinguished from the others in the dense and short pubescence, the vague longitudinal dark stripe running from pron- otum to scutellum, and the structure of male geni- talia. 6. Megacoelum tenuicorne sp. n. (figs. 36-40) Type material. - Holotype male, Canton (23°6'N, 113°12’E), Guangdong Prov. - Paratypes: 12 males, 2 females, with same data as holotype; 1 male, 4 females, Baoxing (30° 18'N, 102°49’E), alt. 950-1360m, Sichuan Prov., 16/18.vi.1963, L. Y. Zheng and H. G. Zou leg.; 1 female, Mt. Emei (29°30’N, 103° 18’E), alt. 600m, Si- chuan Prov., 4.vi.1957, L. Y. Zheng and H. H. Cheng leg.; 1 male (TJNM), Jinchuan (31°27’N, 102°0’E), alt. 2000- 2300m, Sichuan Prov., 10.ix.1963, S. L. Siu leg.; 1 male, 1 female (TJNM), Cujiang (25° 42’N, 102° 0'E), Longsheng County, Guangxi Prov., 24/25.viii.1964, S. L. Liu leg. Description. — Reddish brown, pubescence sparse, elongate and pale. Female larger than male. Clypeus prominent, frons with fine red transverse stripes at lateral sides; antennae thin and brown, segment I with a brown seta on inner margin; rostrum surpassing hind coxae. Collar brown, with a row of long hairs, slightly thicker than the basal diameter of antennal segment II. Pronotum shiny, smooth, posterior lobe indistinctly wrinkled, basal margin slightly raised; calli somewhat reddish. Scu- tellum weakly but densely wrinkled, slightly con- vex. Hemelytra sometimes mottled with some red spots, outer part of corium, embolium and outer part of cuneus semi-transparent, membrane smoky black. Legs brown, apical halves of middle and hind femora, apical 3/4 of middle and hind tibiae red- dish brown, tarsi dark. Thoracic ventral surface dull, brown. Venter red. The knob before left clasper in male distinct (fig. 37). Vesica (fig. 36) with 5 membranous lobes, most of them sha- greened or toothed, one of them with flossy apex. Left clasper elongate, thin and curved (figs. 37-39), right clasper as in fig. 40. Measurements: Body length male 6.46-7.00, fe- male 6.95-7.39; width male 2.11-2.48, female 2.42- 2.60; head apex to cuneal suture male 4.76-5.13, female 5.07-5.39. Head length 1.05-1.09, width 1.14-1.16. Vertex width male 0.36-0.40, female 0.44-0.66. Antennal segment lengths 1.01-1.17 : 2.95-3.55 : 2.39-2.81 : 1.16-1.41. Rostrum length 189 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 3.08-3.62. Pronotum length male 1.12-1.16, female 1.22-1.24; width male 1.84-1.95, female 1.98-2.15. Scutellum length male 0.89-0.96, female 0.92-1.02; width male 0.89-1.02, female 0.99-1.24. Corium length male 3.42-3.49, female 3.72-3.86. Length of anterior femur : tibia : tarsus = 1.84-1.98 : 2.41- 2.52 : 0.63-0.68; length of hind femur : tibia : tarsus = male 2.95-3.02 : 4.59-4.84 : 0.70-0.74, female 3.06-3.18 : 5.03-5.36 : 0.73-0.78. Remarks. - Differs from other species in the slender antennae and the structure of male genita- lia. Figs. 36-45, Megacoelum spp. - 36, Megacoelum tenuicorne sp. n., dorsal view of vesica; 37-39, idem, left clasper; 40, idem, right clasper; 41, Megacoelum yunnananum sp. n., right clasper; 42, 43, idem, right clasper; 44, idem, vesica in dorsal view; 45, idem, vesica, sclerotized broad plate, ventral view. 190 7. Megacoelum yunnananum sp. n. (figs. 41-45) Type material. - Holotype male, Mengla (21°24’N, 101°30’), Yunnan Prov., 24.ix.1979. Description. - Dark brown, with very dense and short pubescence on the hemelytra. Head dull, brown, antennae brown, subbasal part of segment II and basal 1/3 of III yellowish, IV absent. Collar brown, thinner than the basal diameter of antennal segment II. Pronotum black, basal margin nar- rowly yellow and slightly reflexed; calli and the area before and behind dark reddish brown; deeply, densely and transversely wrinkled excepting the smooth calli. Scutellum convex, black, lateral an- gles slightly paler, with fine transverse wrinkles. Hemelytra black with dense and short pubescence, hind part of embolium and outer margin of cuneus reddish, apical 1/3 of cuneus yellowish brown, membrane smoky black. Legs pale brown, apices of anterior and median tibia dark brown. Thoracic ventral surface brown, with black stripes, coxae black. Anterior part of venter reddish brown, pos- terior part black. The knob on pygofer before left clasper in male short (fig. 42). Vesica (fig. 44) with 5 membranous lobes, the ventral surface of the right one sclerotized into a broad plate with its broad knob-like apex bending sideward and armed with numerous minute teeth (fig. 45); the median one with a slender and shagreened (or minutely toothed) shaft and a widened round apex, the latter armed with numerous strong spines directing ven- trad (fig. 44). Left clasper as figs. 42 and 43, right clasper as fig. 41. Measurements: Body length 67.6, width 2.33; head apex to cuneal suture 5.3. Head length 1.02, width 1.16. Vertex width 0.41. Antennal segment lengths = 1.17 : 2.95 : 1.88 : ?. Pronotum length 1.40, width 2.25. Scutellum length 1.07, width 1.15. Corium length 3.52. Length of anterior femur : tibia : tarsus = 1.88 : 2.17 : 0.60; length of posterior femur : tibia : tarsus = 2.30 : 2.84 : 0.61. Remarks. - The narrowly yellow and reflexed pronotal basal margin, deeply and transversely wrinkled disk of pronotum, dense and short heme- lytral pubescence distinctly distinguish this new species from other members of the genus. 8. Megacoelum fuscescens Hsiao, 1963 (figs. 1-5) Megacoelum fuscescens Hsiao, 1963: 440, 442-443, 448; figs. la, 2a, 3. Holotype &: Xishuangbanna-Xiaomen- gyang (22°0’N, 100° 48’E), Yunnan Prov., 24.viii.1958 (in Institute of Zoology, Academia Sinica, Beijing) [examined]. Li « ZHENG: New species of Megacoelum Description. - Dark brown, mottled with lighter hue, with sparse, fine, long and greyish hairs. Head brown, eyes black, frons with several oblique. stripes on both sides. Antennal segment I, most of IV, and apical 1/4 of II dark brown; III, basal 3/4 of II, extreme base of IV yellowish brown, some- times extreme base and middle of II dark brown; I with sparse hairs, II to IV with black setae, II with moderately sparse and long hairs, III and IV with short and dense hairs. Rostrum reaching the apex of hind coxae. Collar opaque, pale yellowish brown or dark brown, slightly thinner than the basal di- ameter of antennal segment II, with a row of long greyish hairs and some pale soft hairs. Pronotum shiny, dark brown, lateral margins and basal part nearly black, extreme basal margin yellowish brown; calli distinctly smooth, reaching antero- lateral angles; posterior pronotal lobe obscurely wrinkled transversely; basal margin slightly curved posterad. Scutellum dark brown, opaque, relatively flat, finely and transversely wrinkled. Hemelytra dark brown, membrane dark. Legs brown, some- times with dark spots; middle femora and tarsus dark, tibial spine shorter than tibial diameter. Tho- racic ventral surface and coxae dull, dark brown. Venter shiny dark brown, sometimes with brown spots, male pygofer with a short knob before left clasper. Vesica with 4 large membranous lobes, among them arised a spicule, slightly flattened and curved dorsally (figs. 1 and 2). Base of left clasper moderately thick, curved and tapered apically (figs. 3 and 4). Right clasper as in fig. 5. Measurements: Body length 6.8-7.4, width 2.2- 2.4; head apex to cuneal suture male 5.2-5.3, female 4.7-4.9. Head length male 1.09, female 1.06-1.07; width male 1.22, female 1.16-1.17. Vertex width male 0.32, female 0.35-0.36. Lengths of antennal segment = male 1.20-1.28 : 2.93-3.15 : 2.67-2.73 : 1.28-1.32, female 1.17-1.19 : 2.68-2.90 : 2.48-2.74 : 1.21-1.29. Rostrum length 3.2. Pronotum length male 1.18-1.43, female 1.12-1.13; width male 1.95- 2.14, female 1.90-1.98. Scutellum length 0.99-1.07, width 1.03-1.15. Corium length 3.43-3.59. Length of anterior femur : tibia : tarsus = 1.98 : 2.36 : 0.62; length of hind femur : tibia : tarsus = 3.22-3.35 : 10325 IOS Remarks. - Hsiao & Meng (1963) remarked that this species is ‘possibly allied to M. straminipes Dist. from India but the first antennal segment shorter, corium concolorously fuscous and apical fourth of posterior femora black’. The species differs from others in the structure of male geni- talia. Material examined. — 1 male, holotype; 2 males, Ruili (24°0’N, 97° 48’E), Yunnan Prov., 28/31.viii.1979, Z. P. Ling and J.X. Cui leg.; 3 males, Lianxian County 191 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 (24° 42’N, 112°18'E), Guangdong Prov., 20/24.ix.1962, L. Y. Zheng and H. H. Cheng leg., on Gramineae and Capsium frutescens L.; 1 male, 2 females, Longzhou County (22°18°N, 106°48’E), Guangxi Prov, 13/18.vii.1964, L. C. Wang leg.; 4 males, 4 females (TJNM), Longzhou County, Guangxi Prov. 15/21.vii.1964, S. L. Liu leg.; 1 female (TJNM), Jianfen- gling (18° 42’N, 108° 48’E), Hainan Prov., 10.v.1964, S. L. Liu leg. 9. Megacoelum clypeale Poppius, 1915 Megacoelum clypeale Poppius, 1915: 14-15. Holotype female: [Taiwan] Kosempo, X, (Deutschen Entomo- logischen Museum, Berlin) [not examined]. 10. Megacoelum minutum Poppius, 1915 Megacoelum minutum Poppius, 1915: 15. Holotype male: [Taiwan], Takao, 18.vii.1907, (Universitats- Museum, Helsingfors) [not examined]. REFERENCES Carvalho, J. C. M., 1959. A catalogue of the Miridae of the world. Part IV. - Arquivos do Museu Nacional, Rio de Janeiro 48: 158-163. Distant, W. L., 1904. The Fauna of British India, including Ceylon and Burma. Rhynchota, 2: 427-430. London. Distant, W. L. , 1910. The Fauna of British India, includ- ing Ceylon and Burma. Rhynchota, 5:235-238. Hsiao, T. Y. & H. L. Meng, 1963: The plant bugs collected from cotton fields in China (Hemiptera-Heteroptera, Miridae). - Acta Zoologica Sinica 15(3): 442-443, 448, figs. la, 2a, 3. 192 Kerzhner, I. M., 1972. New and little known Heteroptera from the Far East of the USSR. - Acta of Zoological . Institute, Academy of Science USSR 52: 276-295. Linnavuori, T., 1963. Contributions to the Miridae fauna of the Far East III. - Annales Entomologici Fennici 29(2): 73-82. Linnavuori, T., 1965. Contributions to the Miridae fauna of the Far East IV. - Annales Entomologici Fennici 31(4): 268-269. Linnavuori, T.,, 1975. Hemiptera of the Sudan, with re- marks of some species of the adjacent countries. 4. Miridae and Isometopidae. - Annales Zoologici Fen- nici 12(1): 1-118. Oshanin, B., 1910. Verzeichnis der Palaearktischen He- mipteren. Band I, Heteroptera. - St. Petersburg. 1087 BE: Poppius, B., 1914. Zur Kenntnis der Miriden, Anthoco- riden und Nabiden Javas und Sumatras. - Tijdschrift voor Entomologie 56 (supplement): 108-111. Poppius, B., 1915. H. Sauter’s Formosa-Ausbeute: Nabi- dae, Anthocoridae, Termatophylidae, Miridae, Isome- topidae und Ceratocombidae (Hemiptera). - Archiv fiir Naturgeschichte 80A (8): 1-80. Wagner, E., 1970/1971. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (Hemiptera Heteroptera). Teil 1. - Entomologische Abhandlungen, Dresden 37 (Supplement): 1-484. Received: 24 February 1991 Revised version accepted: 20 August 1991 DAVID K. MCALPINE Australian Museum, Sydney RELATIONSHIPS OF THE GENUS HETEROCHEILA (DIPTERA: SCIOMYZOIDEA) WITH DESCRIPTION OF A NEW FAMILY McAlpine, D. K., 1991. Relationships of the genus Heterocheila (Diptera: Sciomyzoidea) with description of a new family. - Tijdschrift voor Entomologie 134: 193-199. [ISSN 0040- 7496]. Published 18 December 1991. A morphological comparison is made of the maritime kelp-living genus Heterocheila Rondani with the families of Sciomyzoidea, particularly the Helcomyzidae. It is concluded that it has no particularly close relationship with any one of these families, and the new family Heterocheilidae (halfbridge flies) is established for it. Heteromyza orientalis Macquart, 1843 is a new synonym of Heterocheila buccata (Fallen, 1820). D. K. McAlpine, Australian Museum, Box A285, Sydney South 2000, Australia. Key words. - Diptera; Heterocheilidae, new family; Holarctic. In the course of my recent work on the Coelopi- dae (kelp flies) (D. McAlpine 1991) it became apparent that (1) there is disagreement in recent literature as to the family position of Heterocheila, and (2) it is difficult on morphological grounds to justify the inclusion of Heterocheila in the various families (Coelopidae, Dryomyzidae, Helcomyzi- dae) where it has been recently assigned. For these reasons the present investigation was made. The holarctic genus now known as Heterocheila Rondani, 1857, has often been referred to by the synonymous name Oedoparea Loew, 1862. See both Steyskal (1965), and Gorodkov (1984) for complete synonymy. Included species are: (1) H. buccata (Fallén, 1820) (syn. Heteromyza orientalis Macquart, 1843), from Europe; (2) H. hannai (Cole, 1921) (syn. H. nudiseta Curran, 1933), from Pacific North America. Both species live in stranded kelp (see especially Egglishaw 1960b). Heterocheila (or Oedoparea) has often been placed in the family Helcomyzidae (e. g. Malloch 1933; Hennig 1937, 1973; Steyskal, 1958, 1965; Gorodkov 1984), or in the Dryomyzidae, broadly defined to include Helcomyza Curtis and allied genera (e.g. Czerny 1930; Steyskal 1987; J. McAl- pine 1989). Egglishaw (1960b), Dobson (1976), and Griffiths (1972) have preferred to place Hete- rocheila in the Coelopidae (with some reserva- tions), but I have given definite reasons for its exclusion from the Coelopidae (D. McAlpine 1991), and these need not be repeated here. Useful descriptive and illustrative material of the adult morphology of Heterocheila has been given by Czerny (1930), Hennig (1958), Steyskal (1958, 1962, 1987), and Griffiths (1972). Backlund (1945) and Egglishaw (1960b) have described the egg, larva and puparium of Heterocheila, and the latter also gives important biological information. I have examined a series of adults of both sexes of H. buccata, and collected the species in the field. W. N. Mathis has provided material of H. hannat. The family Helcomyzidae is here considered to include only the genera He/comyza Curtis, Maori- myia Tonnoir & Malloch, and Paractora Bigot. I have given reasons for excluding the group from the Dryomyzidae (D. McAlpine 1991) and sug- gested a closer relationship to the Coelopidae. Use- ful descriptive and illustrative material of the adult morphology of Helcomyzidae has been given in most of the papers mentioned above for Hetero- cheila morphology, also by Malloch (1933). Eggli- shaw (1960a) has described the larva and puparium of Helcomyza, and given biological information. I have examined adult material of all genera and most described species of Helcomyzidae. In the following text I use the superfamily Sci- omyzoidea to include the families Sciomyzidae, Huttoninidae, Helosciomyzidae, Dryomyzidae, Helcomyzidae, Coelopidae, Ropalomeridae, Sepsi- dae, Chamaemyiidae (including Cremifaniidae), Eurychoromyiidae (probably excluding the Gay- omyta complex, see J. McAlpine 1989), and Laux- 193 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 aniidae (including Celyphidae). J. McAlpine (1989) includes the last three families in a separate super- family Lauxanioidea. SIMILARITIES OF HETEROCHEILA TO HELCOMYZIDAE Heterocheila shares with Helcomyzidae s. str. the beach habitat and dependence on stranded kelp, utilising it for larval nutrition as well as shelter. These habits are shared with a number of other Diptera, notably the Coelopidae, the sepsid genus Orygma Meigen, and the sphaerocerid genus (or subgenus) Thoracochaeta Duda. Similarity of hab- its and habitat seems to have caused some conver- gent similarities among these shoreline flies, so that Orygma and Heterocheila have been referred to the Coelopidae in the past. In elevating Helcomyzidae to family status, Mal- loch (1933) emphasised the significance of the sclerotised precoxal bridge, which connects the prosternum to the propleuron on each side. Hen- nig (1958) considered this condition to represent a synapomorphy peculiar to the Helcomyzidae plus Ropalomeridae among the Schizophora, which he therefore considered to constitute a monophyletic group. It is now known that prothoracic precoxal bridges occur in at least some representatives of at least 19 schizophoran families (Speight 1969, and my observations), and many arisals of the condi- tion are acknowledged. Within the superfamily Sci- omyzoidea, the trait occurs also in some sciomyzids as an independent apomorphy. The presence of precoxal bridges is not now usually considered as proof for monophyly of Helcomyzidae and Ropa- lomeridae (Griffiths 1972; J. McAlpine 1989), and additional evidence is needed if monophyly of Heterocheila with Helcomyzidae is to be estab- lished. The precoxal bridge in Heterocheila differs somewhat from that of Helcomyzidae in being markedly narrower. In both Helcomyza and Heterocheila the basal crossvein (crossvein bm-cu or base of M,; accord- ing to divergent applications of the Comstock- Needham notation) lies more obliquely, with ap- proach to a longitudinal orientation, than in Dry- omyza. This condition is unlikely to be a ground- plan condition for Helcomyzidae s. str. in view of the fact that the undoubted helcomyzid Maorimyia has the basal crossvein more nearly transverse than in four available Dryomyza species. I also find some variation in this character in both Coelopidae and Sciomyzidae. It appears to be a rather unsatisfac- tory indicator of phylogenetic affinities in the Sci- omyzoidea. Heterocheila, Maorimyia, and Helcomyza have two spermathecae in the female abdomen, as dis- 194 tinct from the investigated dryomyzids and the apparent groundplan of Coelopidae. On the other hand the number of spermathecae is unrecorded for the helcomyzid genus Paractora. Increasing knowledge of spermathecal numbers in Schizophora has demonstrated that it is a fairly unstable character. The change from three to two spermathecae must have happened many times in the evolution of the Schizophora, if, in fact, the change has always been in the one direction. In the Sciomyzoidea, variation in spermathecal number occurs within the families Huttoninidae, Coelopi- dae, and Sciomyzidae. Other traits shared between Heterocheila and the Helcomyzidae are, so far as I am aware, found in a wide spectrum of sciomyzoid flies and have not been postulated as evidence of closer relationship between these two taxa. Such traits include the development of mollisetae (see D. McAlpine 1991) on various parts of the males, the general brownish grey pruinescent covering of the cuticle, character- istic of many shore-dwelling flies, and the shining parafacial ridges, also found in many flies of mar- itime or sandy habitats. DIFFERENCES BETWEEN HETEROCHEILA AND HELCOMYZIDAE The principal morphological differences be- tween adults of these taxa are given in table 1. They are discussed below in numerical sequence, with the addition of some comments on larval morphol- ogy (18). 1. Hackman & Väisänen (1985) have investi- gated the costal chaetotaxy of the Diptera (includ- ing Heterocheila, op. cit.: fig. 16) and assigned it some taxonomic value at the family and subfamily levels. The additional dorsal and ventral rows of costal setulae present in Heterocheila are in agree- ment with Dryomyzidae and Helosciomyzidae (but not Coelopa frigida (Fabricius) with which Hack- man & Väisänen compare it). The absence of these series in all genera of Helcomyzidae is in contrast to the above taxa. The presence of outstanding anteroventral spines at intervals on the mid region of the costa in all Helcomyzidae separates them sharply from the Dryomyzidae and Heterocheila, and aligns them with Helosciomyzidae and some Coelopidae (e. g. Lopa convexa McAlpine, Gluma keyzeri McAlpine, and Rhis whitleyi McAlpine, see D. McAlpine 1991). 2. The discontinuity of the parafacial suture on its lower part and peculiar angular shining parafa- cial ridge are apparently autapomorphies of the Helcomyzidae. The condition in Heterocheila is probably partly plesiomorphic, though the exten- sive shining ridge may be an autapomorphy. 3. The incised, vertically orientated postgenal fold in Heterocheila, is an unusual feature in the Sciomyzoidea and presumably an autapomorphy. A similar condition occurs in the heleomyzid Nephel- lum dendrophilum (Malloch) (D. McAlpine 1985: Fig. 30). 4. The Helcomyzidae share with the Heloscio- myzidae the narrow median emargination of the face adapted to receive the prelabrum. The emar- gination is absent in Heterocheila and Dryomyzi- dae, but a somewhat similar emargination occurs in some, Coelopidae, though it is doubtful if it is in the groundplan of the latter. Perhaps the median emargination represents a separate apomorphy in each group in which it occurs. 5. The shape of the hypopleural channel in the Helcomyzidae represents a distinctive autapo- morphy, absent in Heterocheila (D. McAlpine 1991: Fig. 9). 6. The significance of this character has been discussed above under ‘Similarities’. 7. The male-restricted apical ventral process of the fore basitarsus is a trait difficult to evaluate phylogenetically, possibly because of irregular loss ina number of lineages (D. McAlpine 1991). In the Sciomyzoidea it is apparently restricted to Helco- myzidae, Coelopidae, and Dryomyzidae, though not uniformly present in the last family. It does not appear to be a synapomorphy as it also occurs in a number of Heleomyzoidea and in Heloclusia im- perfecta Malloch, a somewhat primitive represen- tative of the Nerioidea. This condition seems un- likely to have originated more than once. I therefore regard it as a very ancient trait and it may provide evidence for relationship between Sciomy- zoidea, Heleomyzoidea, and Nerioidea. In this case, its absence in Heterocheila would be a derived state, but not necessarily a synapomorphy with other sciomyzoids in which it is also absent. 8. The phylogenetic significance of the reduction of tergite 6 of the male abdomen in Sciomyzoidea does not appear to have been satisfactorily ex- plained. J. McAlpine (1989) regards reduction of tergite 6 as a groundplan apomorphy of his Scio- myzoidea (as distinct from Lauxanioidea) but in- cludes in Sciomyzoidea such taxa as Heterocheila, Orygma Meigen (Sepsidae), and Rhytidops Lindner (Ropalomeridae) which have a large (and in Orygma, according to J. McAlpine, primarily unreduced) tergite 6. For this and other reasons, I include the lauxanioid families Chamaemyiidae, Lauxaniidae, and Eurychoromyiidae in the Sciomy- zoidea. If evolutionary change in the size of tergite 6 takes place only in the direction of reduction, this might seem to support the idea of a monophyletic group including Dryomyzidae, Helcomyzidae, Coe- MCALPINE: Heterocheila lopidae, and perhaps other families, but excluding Heterocheila, Ropalomeridae, Sepsidae, Chamae- myiidae, Lauxaniidae and Eurychoromyiidae. Con- vergence in this character cannot be excluded, but seems less likely for the complex Dryomyzidae plus Helcomyzidae plus Coelopidae, plus perhaps He- losciomyzidae, as these families show several other somewhat inconsistent similarities. Thus, the size of tergite 6 in Heterocheila makes difficulties for the hypothesis that it is close to the Helcomyzidae or Coelopidae. 9. The aedeagus of Helcomyzidae is more similar to that of Dryomyzidae than to that of Hetero- cheila (Griffiths 1972). My study of the aedeagus in Coelopidae (D. McAlpine 1991) and Heleomyzidae (e. g. D. McAlpine 1967) shows that its structure may be quite unstable above the species level. There are apparently consistent differences in both hypandrium and aedeagus between Hetero- cheila and Helcomyzidae. The hypandrium of Heterocheila has two pairs of processes posteriorly (gonites or parameres) which are absent in helco- myzids. The aedeagus of Heterocheila has a pair of spreading lateral lobes at the junction of the basi- phallus and distiphallus, which is absent in the helcomyzids examined, but it lacks the patch of pubescence near the middle of the length of the distiphallus, which is present in helcomyzids. Griffiths (1972) has emphasised the substantial differences in external male genitalia between Heterocheila and certain helcomyzids. It is possible that these structures should be assigned relatively low reliability as indicators of relationship. To judge from my studies of the Heleomyzidae (D. McAlpine 1967, 1985) and the Coelopidae, the cop- ulatory structures are so unstable above species level that it is difficult to homologise the various processes and lobes across the family. Therefore there is little logic in inferring autapomorphies between special conditions of certain coelopids and those occurring in Heterocheila, which is certainly not so close as a sister group to the Coelopidae. However, I find greater consistency in hypandrial structure within the smaller taxonomic diversity of the Helcomyzidae. 10. The presence of a female-restricted enlarged, isolated, posteriorly directed bristle on each lateral margin of tergites 2 to 4 and sometimes 5 is char- acteristic for most helcomyzids, including species in all 3 genera. It is absent in the Dryomyzidae, but the presence of similar female-restricted bristles in two rather plesiomorphic but not closely related coelopid species (D. McAlpine 1991) may indicate that it was present in the common ancestor of Helcomyzidae and Coelopidae. 11-17. I do not rate these differences in chaeto- taxy very highly, taken individually. Collectively, 195 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Table 1. Differences between Helcomyzidae and Heterocheila (adults). Helcomyzidae 1. Mid regionof costa with spaced anteroventral spines, without continuous series of dorsal and ventral setu- lae. 2. Shining parafacial ridge not continued posteriorly below cheek; parafacial suture obsolete or inter- rupted behind angular section of ridge. 3. Postgenal fold absent. 4. Epistomal margin of face with narrow sinuation to receive prelabrum. 5. Hypopleural channel rather narrow and parallel- sided. 6. Prothoracic precoxal bridge broad, well sclerotised. 7. & fore basitarsus with terminal ventral thumbnail- like process. 8. 6 abdominal tergite 6 much reduced. 9. Aedeagus with densely pubescent zone or zones, without lateral lobes. 10. 9 with one large bristle near each lateral margin on tergites 2-4. 11. Generally only 2 well developed fronto-orbital bris- tles present. 12. Prostigmatal bristle present (sometimes replaced by several long hairs). 13. Median series of acrostichal bristles or setulae not differentiated. 14. Prosternum setulose. 15. Metasternum setulose. 16. Abdominal sternite 1 setulose. 17. © hind tibia with subapical anterior bristle. the seven points indicate a notable degree of div- ergence between the taxa. 18. Larval morphology of Helcomyzidae is only recorded for Helcomyza ustulata Curtis, and it is uncertain if this is representative of the family as a whole. However, in view of the apparently close relationship between the three helcomyzid genera and the very distinctive features of the known larva, these may have some significance for higher classification. According to Egglishaw (1960a, 1960b) the third instar larva of Heterocheila buc- cata has creeping welts of small spines ventrally on 196 Heterocheila Mid region of costa without anteroventral spines, with regular series of dorsal and ventral (as distinct from anterodorsal and anteroventral) setulae. Shining parafacial ridge and associated suture con- tinued below cheek to postgenal region. Postgenal fold distinctly incised. Epistomal margin of face without narrow sinuation. Hypopleural channel expanding rapidly anteriorly. Prothoracic precoxal bridge narrow, not uniformly sclerotised. é (and Q) fore basitarsus without terminal ventral process. 8 abdominal tergite 6 large. Aedeagus without dense pubescence, with pair of lat- eral lobes. Q (and ) without lateral marginal bristles on abdom- inal tergites. Fronto-orbital bristles 3. Prostigmatal bristle absent. Single regular median series of short acrostichal bristles present (in addition to prescutellar pair). Prosternum bare. Metasternum bare. Abdominal sternite 1 bare. Q (and @) hind tibia without subapical anterior bristle segments 3 or 4 to 12 (no such spinose creeping welts present in Helcomyza ustulata, but the cuticle extensively covered with backwardly directed plates), no spine above posterior spiracle (a large spine in this position in Helcomyza), each poste- rior spiracle with two or three groups of short hydrofuge hairs (these absent in Helcomyza). There is also difference in the arrangement and spinose armature of the ridges or processes sur- rounding the anus, and the position of the anus. Egglishaw considers the larvae of Heterocheila and Helcomyza to be so different that they cannot be- long in the same family. I am unable to find in his descriptions and figures any shared distinctive traits, which might provide evidence of relatively close relationship between them, though they share many points of resemblance to numerous other schizophoran larvae. DISCUSSION AND CONCLUSION The few distinctive points of similarity shared between Heterocheila and the family Helcomyzi- dae are unconvincing as indicators of close phy- logenetic relationship. Each of these traits occurs elsewhere in the Sciomyzoidea, and the combina- tion or maritime habitat, a shining section of the parafacial ridge, sclerotised precoxal bridges, and two spermathecae occurs also in the genus Tethina Haliday (Chloropoidea: Tethinidae) though this must be only remotely related to Helcomyzidae. It must be acknowledged that convergence between Heterocheila and Helcomyzidae in these characters is not an improbable event. I have indicated a possible sister-group relation- ship between Coelopidae and Helcomyzidae (D. McAlpine 1991) and this relationship seems to be further supported by costal chaetotaxy. The nu- merous character differences separating Hetero- cheila and Helcomyzidae indicate a marked evolu- tionary divergence, and characters 1, 6, 7,9 and 17 in Table 1 may indicate that Heterocheila is prob- ably not to be included in the monophyletic group Helcomyzidae plus Coelopidae, or even the some- what less surely founded group Dryomyzidae plus Helcomyzidae plus Coelopidae. Heterocheila resembles at least some taxa of the Ropalomeridae in its sclerotised prothoracic prec- oxal bridge, large male tergite 6, and two sper- mathecae. These traits, however, do not provide a stronger case for relationship than those formerly seen to support the now discarded hypothesis of close relationship between Helcomyzidae and Ro- palomeridae. Heterocheila differs from the Ropa- lomeridae inter alia in its less markedly divergent postvertical bristles, differently shaped head cap- sule, unmodified scutellum, absence of setulae on margin of metathoracic spiracle, unmodified femora, and distinct but desclerotised distal section of vein 7. Also Heterocheila lives on north-temper- ate shore-lines, while the ropalomerids are mainly inhabitants of tropical forests of the Americas. Most recent workers (e.g. J. McAlpine 1989) con- sider the Ropalomeridae to be closely related to the Sepsidae. There is no adequate reason for including Heterocheila in this alliance. Heterocheila also resembles the little known neotropical family Eurychoromyiidae in the pro- thoracic precoxal bridge, and the latter has an even MCALPINE: Heterocheila larger male tergite 6 (see J. McAlpine 1968, for morphological details). Heterocheila lacks the fol- lowing distinctive characters of Eurychoromyiidae: body form remarkably stout; chaetotaxy greatly reduced; head structure highly modified; antennal segment 1 (scape) elongate; scutellum abbreviated; mesopleural bristles present; tibiae lacking termi- nal spurs; distal section of vein 7 indistinguishable. There is also considerable difference in the male postabdomens and in other characters. Whereas Heterocheila is restricted to cool temperate shores, eurychoromyiids are only known from Bolivia, a land-locked tropical country. I conclude that the few points of resemblance between these taxa are not indicative of close relationship. Heterocheila is excluded from the Sciomyzidae (perhaps including Phaeomyiinae) because of the prelabrum, and its larvae feed on neither molluscs nor diplopods. Otherwise there are no special points of resemblance and no recent author has suggested a close relationship. Heterocheila is excluded from the Huttoninidae (best regarded as a separate family, not a subfamily of Sciomyzidae or Helosciomyzidae; further dis- cussed in the forthcoming 2nd edition of ‘The In- sects of Australia’) because it lacks the following distinctive features of Huttoninidae: antennal seg- ment 1 without setulae on medial surface; vein 6 abbreviated; distal section of vein 7 scarcely distin- guishable; abdominal sternite 1 vestigial or absent; abdominal tergites 1 and 2 not partly separated by a membranous dorsal line. Huttoninids appear to be mainly forest-living and are endemic to New Zealand. Heterocheila is excluded from the Helosciomy- zidae because it lacks the following apparently con- sistent features of that family: costa with promi- nent spaced anteroventral spines; fronto-orbital plate with at most two bristles; prothoracic pre- coxal bridge absent; abdominal sternite 1 vestigial; abdominal tergite 6 of male much reduced; each posterior spiracle of larva with 4 branched hydro- fuge hairs (Helosciomyza Hendel and Polytocus Lamb). Helosciomyzids are apparently restricted to the south-temperate zone. In my experience they are found in forests, grasslands, swamp-mar- gins, and rocky shores, but are not known to inhabit kelp beds. Heterocheila differs from the families Chamae- myiidae (including Cremifaniidae) and Lauxanii- dae (including Celyphidae) in its complete vein 6 and numerous other traits. These families are mor- phologically more remote from Heterocheila than are most other sciomyzoids (J. McAlpine 1989), they are not associated with kelp, and further com- parison is deemed unnecessary. The exclusion of Heterocheila from all recog- 197 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 nised families of Sciomyzoidea necessitates the set- ting up of a new monogeneric family of this super- family. Heterocheilidae fam. n. Type genus: Heterocheila Rondani, 1857: 104 (present designation). Diagnostic description Actively flying insects of maritime environ- ments; habitus and many structural details typical of Sciomyzoidea; cuticle largely pruinescent and non-shining; parts of thorax, legs, and abdomen with mollisetae, particularly developed in male. Head. - rounded anteriorly; face somewhat con- vex centrally, with shallow fovea below each an- tenna; parafacial ridge shining, extending, to- gether with associated suture, below cheek to postgenal region; cheek deep, setulose; postgenal fold long, distinctly incised, located towards poste- rior surface of head capsule; postvertical bristles often subparellel (slightly variable in orientation); fronto-orbital bristles 3, directed outwards. An- tenna, at rest, subporrect; segment 1 with setulae extending on to medial surface; segment 3 rounded-oval; arista with minute pubescence, seg- ment 5 short, not much exserted; prelabrum mod- erately developed, narrowly separated from face, but not fitting into sinuation on lower margin of latter. Thorax. - Sternopleural suture not ascending posteriorly; hypopleural channel rapidly expand- ing anteriorly; margin of metathoracic spiracle and sternopleuron bare; prosternum broadly triangu- lar, with narrowly sclerotised precoxal bridges; other significant characters of chaetotaxy and ves- titure as in Table 1. Each tibia with one preapical dorsal bristle; all basitarsi without terminal ventral process; last 2 tarsal segments depressed and dis- tally expanded. Costa unbroken, extending to vein 4, beyond end of subcosta with an anterodorsal series of closely placed spinules and a regular series each of dorsal, anteroventral, and ventral setulae or hairs, without spaced anterior or anteroventral spines; vein 6 visible approximately to wing mar- gin; vein 7 beyond alula represented by long, curved crease in membrane. Abdomen. - Tergites 1 and 2 incompletely se- parated by a dorsal transverse membranous line; sternite 1 short, but rather well sclerotised, bare; 7 pairs of abdominal spiracles situated in pleural membrane. Male postabdomen: tergite 6 shorter than preceding tergites but not greatly reduced, almost symmetrical, setulose; sternites 6 and 7 placed on left side; tergites 7 and 8 absent or not definitely identifiable (‘8T’ of Steyskal is the scle- 198 rite generally understood to be the inverted ster- nite 8); surstylus articulated basally, with basal an- terior expansion or secondary lobe; cerci separate; basiphallus with broadly expanded membranous posterior lobe; distiphallus with complex scleroti- sation and no pubescence. Female postabdomen only moderately extensile, with all segments short; tergites and sternites 6-8, epiproct, and hypoproct all well sclerotised and setulose; cerci separate, el- ongate. Nomenclatural notes I have examined the holotype of Heteromyza orientalis Macquart, 1843, in the National Museum of Natural History, Paris, and find it to be identical with Heterocheila buccata (Fallen, 1820) (syn. n.). The given type locality of Macquart’s species, Java, is erroneous; the species is probably restricted to Europe, where it is apparently absent from warmer, southern areas. I propose the common names half-bridge flies, for representatives of the family Heterocheilidae, and bridge flies for representatives of the family Helcomyzidae. These names refer to the develop- ment of the prothoracic precoxal bridge. ACKNOWLEDGEMENTS I am indebted to M. C. D. Speight for help in finding Heterocheila in the field, to L. Tsacas and M. Baylac for the opportunity to examine type material in the National Museum of Natural His- tory, Paris, and to W. N. Mathis and J. R. Vockeroth for study material. REFERENCES Backlund, H. O, 1945. Larvae and pupae of Heterochila buccata Fall. compared with the supposed larva of Helcomyza ustulata Curtis (Dipt., Dryomyzidae). - Kungliga Fysiografiska Sallskapets i Lund Förhand- linger 15(6): 5 pp. Czerny, L., 1930. Dryomyzidae. - In: Lindner, E. (editor). Die Fliegen der Palaearktischen Region 38a: 1-8. E. Schweizerbart’sche Verlagbuchhandlung, Stuttgart. Dobson, T., 1976. Chapter 16. Seaweed flies (Diptera: Coelopidae, etc.). — In: Cheng, L. (editor). Marine insects: 447-463. North-Holland Publishing Com- pany, Amsterdam. Egglishaw, H. J., 1960a. The life-history of Helcomyza ustulata Curt. (Dipt., Dryomyzidae). - Entomologist’s monthly Magazine 96: 39-42. Egglishaw, H. J., 1960b. Studies on the family Coelopidae (Diptera). - Transactions of the Royal Entomological Society 112: 109-140. Gorodkov, K. B., 1984. Family Helcomyzidae. - In: Sods, A. & L. Papp (editors). Catalogue of Palaearctic Dip- tera 9: 149-150. Elsevier, Amsterdam etc. Griffiths, G. C. D., 1972. The phylogenetic classification of Diptera Cyclorrhapha with special reference to the structure of the male postabdomen. — W. Junk, The Hague. Hackman, W. & R. Väisänen, 1985. The evolution and phylogenetic significance of the costal chaetotaxy in the Diptera. - Annales Zoologicae Fennicae 22: 169- 203. Hennig, W., 1937. Coelopidae. - In: Lindner, E. (editor). Die Fliegen der Palaearktischen Region 52: 39pp. E. Schweizerbart'sche Verlagbuchhandlund, Stuttgart. Hennig, W., 1958. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehun- gen. - Beiträge zur Entomologie 8: 505-688. Hennig, W., 1973. 31. Diptera (Zweiflügler). - Handbuch der Zoologie 4(2) 2: 337 + 4 unnumbered pp. Malloch, J. R., 1933. Acalyptrata (Helomyzidae, Trypeti- dae, Sciomyzidae, Sapromyzidae, etc.). - Diptera of Patagonia and South Chile 6: 177-391, pl. 2-6. McAlpine, D. K., 1967. The Australian species of Diplo- geomyza and allied genera (Diptera, Heleomyzidae). - Proceedings of the Linnean Society of New South Wales 92: 74-106. McAlpine, D. K., 1985. The Australian genera of Heleo- myzidae (Diptera: Schizophora) and a reclassification of the family into tribes. - Records of the Australian Museum 36: 203-251. McAlpine, D. K., 1991. Review of the Australian kelp flies (Diptera: Coelopidae). — Systematic Entomology 16: 29-84. McAlpine, J. F, 1968. Taxonomic notes on Eurychoro- myta mallea (Diptera: Eurychoromyiidae). - The Can- adian Entomologist 100: 819-823. MCALPINE: Heterocheila McAlpine, J. F, 1989. Phylogeny and classification of the Muscomorpha. - In: McAlpine, J.F. (editor). Manual of Neartic Diptera 3: 1397-1518. - Canadian Govern- ment Publishing Centre, Hull, Quebec. Rondani, C., 1857. Dipterologiae Italicae prodromus. Vol. 3: Species Italicae ordinis dipterorum in genera cha- racteribus definita, ordinatim collectae, methodo ana- litica distinctae, et novis vel minus cognitis descriptis, Pars prima: Oestridae, Syrpfhidae [sic], Conopidae, 264 pp., 1 fig. - Parmae [= Parma]. Speight, M. C. D., 1969. The prothoracic morphology of acalyptrates (Diptera) and its use in systematics. — Transactions of the Royal Entomological Society 121: 325-421. Steyskal, G. C., 1958. Notes on nearctic Helcomyzidae and Dryomyzidae (Diptera Acalyptratae). - Papers of the Michigan Academy of Science, Arts, and Letters 43: 133-143. Steyskal, G. C., 1962. Notes on palaearctic Dryomyzidae, Helcomyzidae and Sciomyzidae (Diptera). - Notulae Entomologicae 42: 71-72. Steyskal, G. C., 1965. Helcomyzidae. - In: Stone, A. et al. (editors). A Catalog of the Diptera of America north of Mexico: 678-679. - United States Department of Agriculture, Washington DC. Steyskal, G. C., 1987. Dryomyzidae. - In: McAlpine, J. E (editor). Manual of Nearctic Diptera 2: 923-926. — Canadian Government Publishing Centre, Hull, Qu- ebec. Received: 4 June 1991 Accepted: 25 July 1991 199 BOOK REVIEW Knight, W.J. & J. D. Holloway (Eds), 1990. Insects and the rain forests of South East Asia (Wallacea). - The Entomological Society of London, London: p. i-iv + 1- 343, figs. [ISBN 0 90546 76 3. Price £ 33.50 (incl. p & p)]. This publication is one of the results of ‘Project Wallace’ and includes the papers presented at a special symposium to highlight some of the major research topics undertaken during that expedition. Readers of the Tijdschrift voor Entomologie cer- tainly have heard about this one year-long expedi- tion to North Sulawesi (1985), since various results have been published in earlier volumes of this journal. : As one of the participants myself, I am still impressed by the efforts the Royal Entomological Society of London (RESL) has made for the benefit of entomology, commemorating the society’s 150th anniversary. Anybody who has any expe- rience in getting things organized for his own trips, or perhaps for smaller expeditions, in tropical countries, knows about the difficulties. Logistics under primitive conditions and co-ordination of enforced co-operation between scientists were some of the organizational pitfalls the leaders of this expedition had to observe. Also the Indonesian government should be congratulated with usually adequate handling their problems. Dozens of ser- vicemen and scientists, flying in and out via Jakarta, all had to get their correct ‘surat’ (letters) on time. Especially the bureau of Lembaga Ilmu Pengeta- huan Indonesia, co-ordinating all the foreign scien- tific activities in Indonesia, must have heaved a sigh of releave on December 31st, 1985. From a biogeographical point of view, Sulawesi is one of the most interesting places on earth, as was already noticed by Alfred Russel Wallace. The geology of Sulawesi is extremely complex. The expedition was held in the Dumoga-Bone National Park, which is approximately central in the Min- ahasa peninsula. This peninsula is considered to be a part of an island of Oriental derivation. The central-eastern and south-eastern peninsula to- gether formed once another island, probably of Australasian origin. Both islands collided only in the Miocene, and the latter island emerged above sea level only since that time. The National Park is fully covered with rain forest, and the protection of the forest is of high importance for the safe- guarding of the water catchment of the Dumoga Plain, which is now a transmigration area. Although the present volume has to be consi- dered as the final report of the expedition, it is apparent from most papers that the study of the 200 material has only just begun. The insect fauna of Sulawesi was nearly completely unknown, except for some groups attracting also the general collec- tor. With so many specialists collecting their own groups, and the extensive general sampling pro- grams with malaise traps, light traps and fogging, literally millions of specimens have found their way to museums. It will take long until all the specimens are named. Just a few examples. In the Ephemeroptera not one species was reported from the whole of Sulawesi; now Edmunds and Polhe- mus mention the existence of twenty genera with one to four undescribed species per genus. Prelimi- nary sorting of the beetles collected by the British Museum (Natural History) revealed an estimated total of more than 5500 species. For the general reader interested in biodiversity and ecology, Hammond’s report on the BMNH’s study program, including large scale malaise trap- ping and fogging (a method to sample the canopy fauna using insectides), is highly relevant, both for the description of the methods as well as for the results. For instance, it is claimed now that the percentage of plant-associated beetles in tropical ecosystems may prove to be far less than an esti- mation based on described species alone would suggest. In the present studies, no less than 29% of the beetles collected were predators and only 17% herbivores. Hammond also concludes that all but 18 of the 130 beetle family-groups collected in northern Sulawesi were found at light. For 49 bee- tle groups light trapping produced even a greater number of species than any other method. For biogeographers there is also a lot of new information. Although the fauna of Sulawesi seems to be Oriental rather than Australian, a direct fau- nal exchange between Sulawesi and Borneo seems to have occurred very rarely. Vane-Wright presents good evidence that the relationship between the fauna of the Philippines and Sulawesi is stronger than generally accepted. Also, it seems that Su- lawesi has been an important source for the Moluc- cas plus New Guinea. A sister relation between Sulawesi and eastern Indonesia has been confirmed in several taxonomically unrelated groups (e. g. the danaid genus Idea, Cicadidae). Of course, Sulawesi is particularly of interest for its huge percentage of endemics, as can be expected for an isolated area that increased considerably in size after a long period of relative stability. Apart from the taxo- nomic groups mentioned before, there are also contributions on Hesperiidae, rodent ectoparas- ites, aquatic Heteroptera, Phoridae, Tetrigidae and catantopine Acrididae. Furthermore, there are papers on the physiography and vegetation of the Continuation on p. 278 ERIK J. van NIEUKERKEN! & RIMANTAS PUPLESIS? INational Museum of Natural History, Leiden *Zoologijos Katedra VPI, Vilnius TAXONOMY AND DISTRIBUTION OF THE TRIFURCULA (GLAUCOLEPIS) RAIKHONAE GROUP (LEPIDOPTERA: NEPTICULIDAE) Nieukerken, E. J. van & R. Puplesis, 1991. Taxonomy and distribution of the Trifurcula (Glaucolepis) raikhonae group (Lepidoptera: Nepticulidae). - Tijdschrift voor Entomo- logie, 134: 201-210, figs. 1-22. [1ssw 0040-7496]. Published 18 December 1991. Trifurcula (Glaucolepis) melanoptera sp. n. is described from southern Europe, where it is widespread. It is closely related to the Central Asian T. (Glaucolepis) raikhonae Puplesis, which is redescribed. Together they form the raikhonae species group. Sinop- ticula Yang is here synonymized with Glaucolepis Braun, its only species S. sinica Yang is tentatively regarded as closely related, if not conspecific with T. ratkhonae. The distribution is mapped. On the basis of recent Chinese findings the larvae are assumed to be gall-makers on Prunus branches. Some phylogenetic and biogeographic remarks are given. Correspondence: E. J. van Nieukerken, National Museum of Natural History, Postbus 9517, NL-2300 RA Leiden, The Netherlands. Key-words. - Microlepidoptera, Central Asia, Southern Europe, biogeography, phylo- geny, gall-former, new species. The subgenus Glaucolepis Braun, 1917, is one of three subgenera of Trifurcula Zeller, 1848 (van Nieukerken 1986b, 1990). It comprises to date 21 described species (see van Nieukerken 1986a, 1986b) and a large number of undescribed species. Most of these occur in the Mediterranean region. The type species T. saccharella (Braun, 1912) is the only Nearctic species. Further, one undescribed species occurs in Japan, one in North-East India and T. raikhonae (Puplesis, 1985) in Central Asia. In treating Glaucolepis as a subgenus, we follow the opinion of the senior author, whereas the junior author prefers to treat it as separate genus (Puple- sis 1985). Amongst unidentified material from southern Europe, one apparently widespread species very similar to T. raikhonae was found by both authors independently. Although a revision of the many mediterranean species is not yet practicable at this stage, we would like to single out this peculiar species, because it forms a well defined monophy- letic group with T. raikhonae: the raikhonae spe- cies group. This group can easily be distinguished from all other species in the subgenus, both on externals and genitalia and possibly also on biol- ogy. It was also desirable to have the new name available for a general work on Nepticulidae of the Soviet-Union and other faunistical reports. We further take the opportunity to redescribe T. raik- honae for the first time in English, including new data and the description of the female, and to dis- cuss a recently described Chinese taxon. Some re- marks on phylogeny and biogeography conclude this paper. MATERIAL AND METHODS Genitalia were prepared as described by van Nieukerken et al. (1990). Line figures of genitalia were prepared with a Zeiss Axioskop with drawing apparatus, both from genitalia in glycerin and per- manent mounts. SEM micrographs were taken with a Jeol JSM 840A scanning electron micro- scope. Specimens were air-dried, mounted on stubs and gold-coated. Measurements of genitalia are taken with a Zeiss Axioskop at 200X, and are accurate at the nearest 5 um. Capsule length is measured along mid-line, from tip of tegumen (pseuduncus) to anterior mar- gin of vinculum, in middle. All measurements based on sample size of at least five specimens are accompanied by mean, standard deviation and sam- ple size in brackets. 201 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Locality names are spelled in accordance with the Times Atlas of the World (Comprehensive edition 1975 and later). For all European localities the appropriate UTM grid references are given, see also van Nieukerken (1990). Abbreviations (codens) for depositories follow Arnett & Samuelson (1986), with the addition of ZKVWV (Zoologijos Katedra VPI, Vilnius, Lithuania). SYSTEMATIC PART Subgenus Glaucolepis Braun Glaucolepis Braun, 1917. Fedalmia Beirne, 1945. Trifurcula (Glaucolepis); van Nieukerken 1986b: 65. Sinopticula Yang, 1989: 79, 81. Type-species: Sinopticula sinica Yang, 1989 (by monotypy and original designa- tion). Syn. n. Recently, Yang (1989) described Sinopticula sin- ica from China. Although we have been unable to borrow material, we are convinced on the basis of description and figures that this species is closely related to, if not the same as T. raikbonae. We therefore synonymize Sinopticula here with Glau- 202 Figs. 1-3. Trifurcula raikhonae group, males. - 1, T. rat- khonae, Dushanbe, underside wings; 2, T. melanoptera, Krk, underside wings; 3, T.melanoptera, Krk, habitus. colepis, and hence with Trifurcula. For description of genus and subgenus and detailed references we refer to van Nieukerken (1986b). The Trifurcula raikhonae group This group is established here to accommodate T. raikhonae, T. sinica and T. melanoptera sp. n. It can be distinguished from other species of Trifur- cula (Glaucolepis) by the absence of the ‘velvet’ patch of raised scales on the male hindwing under- side, the presence of dark brown or black androcon- ial scales on the forewing underside, the absence of rows of spines near phallotrema in the aedeagus and the relatively well developed accessory sac in the female genitalia. Trifurcula (Glaucolepis) raikhonae (Puplesis) (figs. 1, 3, 5-7, 14, 17-19) Glaucolepis raikhonae Puplesis, 1985: 71. Holotype 6: USSR, Tadzhikistan, about 30 km S Dushanbe, Kon- dara canyon, 10-20 June 1982, Sherniyazova (ZMAS) [examined ]. Trifurcula raikhonae; van Nieukerken 1986b: 68. VAN NIEUKERKEN & PUPLESIS: Trifurcula raikhonae group Fig. 4. Distribution of Trifurcula raikhonae group in east Palaearctic. -Circles: T. raikhonae, square: T. sinica (Xi'an), triangle: easternmost record of T. melanoptera (Kopet Dag). Description Male (fig. 1). - Forewing length 3.1-3.9 mm (3.42 + 0.19, n=32). Head: frontal tuft pale orange to ferruginous, collar slightly paler. Antenna brown, with 45-56 segments (50.3 + 3.0, n=19); scape yellowish white. Thorax fuscous. Forewings fuscous, slightly irrorate with white because of paler scale bases; dorsum often paler, with some white scales at tornus, sometimes extending further; cilia-line more or less distinct, terminal cilia pale grey. Forewing underside anterior of fold usually with an elongate patch of about % wing- length and % wing-width, with dark fuscous to black androconial scales (fig. 1), usually with blue iridescence, occasionally patch absent or paler and almost invisible. Hindwing grey, humeral lobe with a small group of extremely small whitish special scales, macroscopically hardly visible, dis- tinct under SEM (figs. 17-19): they are strongly ribbed, with small holes between the ribs; in fig. 19 it seems that the scale has an apical pore. Costal bristles normal; underside without velvet patch as most other Trifurcula (see van Nieukerken 1986b, 1990). Abdomen grey-brown dorsally, yellowish ventrally; three pairs of anal tufts yellowish grey. Female. - Forewing length 3.2-4.0 mm (3.56 + 0.26, n=8). Antenna with 42-49 segments (44.8 + 2.6, n=5). Forewing underside grey-brown, further as male. Male genitalia (figs. 5-7). - Capsule length 390- 470 um (430.0 + 27.7, n=6). Vinculum with very long, truncate, anterior extension, length (190) 245-280 um, ca (0.5) 0.6X capsule length. Tegu- men forming a broadly truncate pseuduncus. Uncus with slightly widened tip, truncate, lateral arms broadly shouldered, very conspicuous. Gnathos with broadly rounded central element, lateral arms inserted on vinculum near valva base. Valva short, 165-185 um long, almost triangular, with pointed narrow tip of about Ys valva length, inner margin with slight bulges in second half; sublateral process relatively short, transverse bar of transtilla not sclerotized, but present. Aedeagus 365-430 um (400.0 + 22.4, n=6) long, with asymmetrical ven- tral lobe, more sclerotized at left side; single long cornutus 185-235 um long, about % aedeagus length, slightly curved, tip more or less rounded; cathrema large, almost triangular, no additional cornuti. Female genitalia (fig. 14). - Terminal segments blunt and rounded; T8 with groups of many scales and several setae, anal papillae with 34-53 setae each. Vestibulum with conspicuous folded acces- sory sac. Ductus spermathecae outer canal wide and distinct, inner canal strongly sclerotized, with long basal straight part, followed by 7 convolutions. Bursa relatively small, covered with transverse rows of pectinations and single spicules; reticulate signa very inconspicuous, only visible under high magnification, using phase-contrast or differential interference contrast; cells mostly incomplete, rec- ognized by longitudinal groups of pectinations. Diagnosis From other species of Glaucolepis, raikhonae differs by the absence of a velvet patch on the male 203 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 5-13. Trifurcula raikhonae group, male genitalia - 5-7, T. raikhonae, Tadzhikistan, slide EJvN 2787; 8-13, T. melanoptera: 9-11, holotype, slide EJvN 2786; 8, 12, Turkey, slide EJvN 2784; 13, Armeniya, slide RP. - 5, 8, 9, capsule, ventral aspect; 6, 10, 12, left valva, dorsal aspect; 7, 11, aedeagus, ventral aspect; 13, cornutus, ventral apsect, but slightly rotated compared to fig. 11. Scales: 0.1. mm, top to raikhonae, bottom to melanoptera. 204 hindwing underside, the relatively large size and the frequently present androconial patch on fore- wing underside. It is currently the only known Central Asiatic Glaucolepis, and can therefore hardly be confused with any other nepticulid of the area. It differs from the only other Central Asiatic Trifurcula species, T. puplesisi van Nieukerken, 1990, by its larger size, dark colour and in male by absence of yellow patch on forewing underside and velvet patch on hindwing. For differences with melanoptera see below. Biology Unknown, but considering the biology of the closely related, if not conspecific sinica, possibly also a gall-maker of Prunus. Various Prunus species are widespread and common in the Central Asian mountains. Adults found from May to August, at light. Distribution (fig. 4) Widespread in Central Asian mountains: west- ern and central Tyan Shan (Kazakhstan and Kir- giziya), Gissarskiy (Hissar) ridge (Tadzhikistan), and northern Kugitangtau mountains (Uzbekis- tan) and in central Afghanistan. Previously only known from two specimens in Tadzhikistan and Kirgiziya (Puplesis 1985). Not found in lowland desert areas. Material examined. - Afghanistan: 1 8, Paghman, 30 km NW Kabul, 2100 m, 20-30.vii.1962, E. & A. Vartian (NHMW); 1 8, same locality, 2500 m, 15-18.vii.1965, Kasy & Vartian (NHMW).- USSR: Kazakhstan: 11 @, 1 9, Tyan Shan, 90 km E Chimkent, 1300 m, Aksu Dzhabagly, 7- 12.viii.1987, Sheriyazova (ZKVV, RMNH) - Kirgiziya: 1 & (paratype), 5 km SW Naryn, 26.vii.1981, Sinev (ZMAS); 2 8, 2 9, Sosnovka, 40 km S Kara Balty, 13.viii.1987, Lvovskyi (ZMAS). — Uzbekistan: 3 @, env. Derbent, 18.v.1985, R. Puplesis (ZKVV, RMNH). - Tadzhikistan: ca 90 4, 9, 30 km N Dushanbe, Kondara, 27.vi-20.viii.1986, R. Puplesis (ZKVV, RMNH); 13 @, 6 9, idem, but 17- 20.viii.1989 (ZKVV). Trifurcula (Glaucolepis) sinica (Yang) comb. n. Sinopticula sinica Yang, 1989: 80, 82. Holotype @: China, Shaanxi prov., Xi'an, emerged 24-30.iv.1985, from galls on Prunus, Yan-wen (Beijing Agricultural Uni- versity) [not examined] Remarks The relatively detailed description of Sinopticula sinica Yang, 1989 and the small, but distinct figures show many similarities to raikhonae. The slight differences in the form of the valvae in Yang's figure from the usual shape in raikhonae might be VAN NIEUKERKEN & PUPLESIS: Trifurcula raikhonae group due to the preparation technique. Without detailed examination we would rather not synonymize sın- ica here, although we suspect that it indeed is con- specific with raskhonae. The locality Xi'an is at the east end of the almost continuous Central Asian mountain ranges, in which ratkhonae presumably is widespread. Biology According to Yang (1989) the larvae make galls in young branches of Prunus cerasifera Ehrh., P. dulcis (Miller) D. A. Webb (= P. amygdalus Bartsch) and P. persica (L.) Batsch. It is not clear from the description, how these galls look like. Adults emerged in April. Distribution (fig. 4) Only known from China, Shaanxi province. Trifurcula (Glaucolepis) melanoptera sp. n. (figs. 2, 3, 8-13, 15, 16, 20-22). Type-material. - Holotype: 8, Yugoslavia (Croatia), Krk, Misucaynica [UTM: 33T VK78], 4.viii.1986, G. Bal- dizzone, Genitalia slide E. J. van Nieukerken No. 2786 (RMNH). — Paratypes: 52 4, 3 9: Austria: 1 6, Hackels- berg, N. Neusiedlersee (N. Burgenland) [UTM: 33U NV52], 24.viii.1973, F. Kasy (NHMw). - Czechoslovakia: 3 6, Slovakia, Turfia n. B. [UTM: 34U DU9183], 4- S.viii.1990, A. Laëtüvka (coll. Laëtüvka); 1 4, Slovakia, Tinianska stran [NE Michalovce] [UTM: 34U EV60], 20.viii.1989, Z. Tekär. - France: 1 9, Les Mees (Alpes de Haute Provence) [UTM 31T GJ37], 28.viii.1985, G. Lan- gohr (RMNH); 1 4, Chapeau [not traced, near Digne?], 27.vii.1903, Chrétien (MNHN); 12 6, 1 9, Viens (Vau- cluse) [UTM: 31T GJ06], 9.viii.1973, 14.vii-23.viii.1974, 6.viii.1975, 9-27.viii.1976, 15.viii.1979, 8.viii.1980, 12.viii.1982, R. Buvat (RMNH, coll. Buvat). - Italy: 1 &, Sardinia, Bacu Trotu, Ortuabis 800 m [uTM: 32S NK11], 23.viii.1978, Gg. Derra (coll. Derra).- Spain: 1 6, Cadalso (Madrid) [uTM: 30T UK86], 15.vii.1985, C. Gielis (coll. Gielis); 2 6, Cadalso de los Vidrios, 2 km E (Madrid), [UTM: 30T] UK8062, 7.viii.1986, at light ML, mattoral, cult. area, E. J. van Nieukerken & S. Richter (RMNH); 1 6, Noguera (Teruel), [UTM: 30T XK17], 9-10.viii.1989, C. Gielis (coll. Gielis); 1 @, Paterna del Madera (Alba- cete), 1350 m [UTM: 30S WH57], 18.vii.1986. C. Gielis (RMNH); 1 9, Pto de Mora (Granada) 1350 m [UTM: 30S VG52] 22.vii.1986, C. Gielis (RMNH); 1 8, Riazza (Sego- via), [UTM: 30T VL57], 3.viii.1986, C. Gielis (coll. Gielis); 1 6, San Miguel de Valero (Salamanca), 3 km S Linares de Riofrio, [UTM: 30T] TKS9, 2.viii.1986, at light ML, Quercus pyrenaica forest and heathland, 850 m, E. J. van Nieukerken & S. Richter (RMNH); 2 8, Vega del Codorno (Cuenca), 1350 m, [UTM: 30T WK97], 23.vii.1985, at light, J. H. Kuchlein (RMNH, coll. Kuchlein). - USSR, Turkmeniya: 1 6, 30 km E Kara Kala, (Western Kopet Dag range), env. Juvankala [UTM: 40S DH35], 18.viii.1988, R. Puplesis (RMNH) — USSR, Ukraina: 2 3, Crimea, Kara Dag, 20 km W Feodosia [UTM: 36T XQ78], 205 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 14, 15. Female genitalia, dorsal aspect. - 14, T. raikhonae, Tadzhikistan, slide EJvN 3205; 15, T. melanoptera, paratype, Spain, slide EJvN 3102. Scale: 0.2 mm. 3.viii.1986, Buhashkim (ZMAS); 1 d, same data, 13.vii.1987, Sinev (ZMAS); 6 À, same data, 15-22.vii.1987, R. Puplesis (ZKVV, RMNH). - Yugoslavia (Croatia): 1 4, Krk, no further data, 10.viii.1975, at light, G. Baldizzone (Coll. Baldizzone); 2 6, 1 9, Krk, Draga Baska [UTM: 33T VK78], 30.vii.1986, 15.viii.1988, G. Baldizzone (RMNH, coll. Baldizzone); 2 &, Krk, Misucaynica [UTM: 33T VK78], 19.viii.1986, G. Baldizzone (coll. Baldizzone); 10 @, Krk, road Krk-Vrbnik [UTM: 33T VK78], 2, 18.viii.1987, 20.vii-11.viii.1988, G. Baldizzone (RMNH, ZKVV, coll. Baldizzone). Material excluded from type series (34 &). - Hungary: 30 & (all in poor condition), Budapest, Julianna Major, apple orchard, 9+11.viii.1991, glued pheromone traps, M. Tóth (RMNH). — Italy: 1 g, Latina, Monti Aurunci, 850 m, 5 km N Itri [UTM: 33T UF77], 4-11.viii.1972, R. Johansson (coll. Johansson). - Turkey: 2 &, 10 km NW Kizilcahaman (Ankara), 1150-1250 m [UTM: 36T VK68], 6-7.viii.1989, Fibiger & Esser (ZMUC). - USSR, Arme- niya: 1 g, Chosrov reserve [UTM: 38S MK82], 20.ix.1986, P. Ivinskis (ZKVV). Description Male (figs. 2, 3). - Forewing length 2.4-3.0 mm (2.74 + 0.12, 29), wingspan 5.8-6.9 mm. Head: frontal tuft pale orange to ferruginous, collar 206 slightly paler. Antenna brown, with 40-45 (42.2 + 1.4, 23) segments; scape yellowish white. Thorax fuscous, often distally paler. Forewings fuscous, slightly irrorate with white because of paler scale bases; dorsum with narrow stripe of white scales, occasionally reduced to white tornal spot; cilia-line more or less distinct, terminal cilia pale grey. Fore- wing underside anterior of fold almost completely covered with dark fuscous to black androconial scales, usually with blue iridescence, except at wing tip, which is greyish-brown. Ultrastructurally with many circular holes between ribs (figs. 21, 22). Hindwing grey, humeral lobe with few fuscous androconial scales, as forewing; occasionally these scales extending on hindwing upperside along Rs+M, forming an elongate patch; costal bristles forming a short brown hair-pencil; near frenulum a group of microtrichia, no scales (fig. 20); under- side without velvet patch as most other Trifurcula. Abdomen grey-brown dorsally, yellowish ven- trally; three pairs of anal tufts yellowish grey. Female. - Forewing length 2.75-3.0 mm, wing- span 6.4-6.7 mm. Antenna with + 39 segments. Forewing underside dark grey-brown, darker than VAN NIEUKERKEN & PUPLESIS: Trifurcula raikhonae group Fig. 16. Distribution of Trifurcula melanoptera, mapped on 50 km squares of UTM grid. hindwing. Otherwise as male. Ovipositor wide and truncate. Male genitalia (figs. 8-13). - Capsule length 305- 390 um (345 + 20.3, n=11). Vinculum with very long, truncate, anterior extension, length 200-260 um, ca 0.6-0.7X capsule length. Tegumen forming a broadly truncate pseuduncus. Uncus with slightly widened tip, or pointed, lateral arms shouldered, but less than in razkhonae. Gnathos with broadly rounded central element, lateral arms inserted on vinculum near valva base. Valva short, 140-165 um (149.1 + 6.3, n=11) long, almost triangular, with pointed narrow tip of about % to % valva length, inner margin with slight bulges in second half; sublateral process relatively short, transverse bar of transtilla not sclerotized. Aedeagus 325-390 um (356.8 + 18.4, n=11) long, with distinct asymmet- rical ventral lobe, sclerotized at left side, with an- terior point; single long cornutus 130-205 um long, slightly less than % aedeagus length, curved, tip distinctly bifurcate, with rounded lobes forming an angle of almost 180° (fig. 13); several additional small cornuti, some forming ‘pectinations’; ca- threma large, almost triangular. Female genitalia (fig. 15). - Very similar to those of T. ratkhonae, but smaller. Anal papillae with 25- 30 setae each. Accessory sac of bursa even larger than in raskhonae. Ductus spermathecae with 5% to 6/2 convolutions. Diagnosis Very similar to T. ratkhonae, but distinctly smaller and in male with much larger androconial patch on forewing underside and with small sim- ilar patch on humeral lobe of hindwing. In male genitalia easily distinguished by the bifurcate cor- nutus. In female genitalia by larger accessory sac and smaller bursa. From other southern European species of Trifurcula, males of melanoptera differ by the absence of a velvet patch on the male hind- wing underside, and the large black or fuscous an- droconial patch on the forewing underside. Variability Specimens from Turkey and Armenyia differ from the remaining material by the presence of androconial scales on the hindwing upperside. Also the genitalia of especially the Armenyian specimen differ slightly in size, shape and position of cornu- tus. However, these characters seem to vary within the whole distribution area, with the Armenyian specimen as the extreme example. Since we also observed few androconial scales on the hindwings of some other specimens, we tentatively assume that melanoptera forms one widespread, slightly variable species, until more data become available. It should also be noted that the specimen from Italy, Itri, has relatively small genitalia, which look slightly different, but tentatively is regarded as an aberration. Both this specimen and the Turkish and Armenyian specimens are excluded from the type series. Biology Immature stages and hostplant unknown. Judg- ing from its close relationships with T. raikhonae (and hence sinica), it is not impossible that also melanoptera is a gall-maker on Prunus. In some localities, visited by the senior author, such as Viens, France, Prunus spinosa L. was abundant; galls were, however, not yet seen. The record of 207 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 17-22. Trifurcula raikhonae group, male scale structures, scanning micrographs. - 17-19, T. ratkhonae, hindwing humeral lobe, upperside: patch of small special scales, with details. 20-22, T. melanoptera: 20, hindwing humeral lobe with group of microtrichia and special scales (left); 21, 22, androconial scales of forewing underside with many circular holes. Scale bars: 100 um (17), 10 um (18, 20, 21), 1 um (19, 22). males taken in pheromone traps in an apple or- chard in Budapest does not contradict this, since this orchard contains several fruit trees, including Prunus spp., and is surrounded by shrub with much Prunus spinosa (G. Szöcs in litt.). Adults caught from 13 July to 28 August, in Armeniya on 20 September, most likely univoltine. Distribution (fig. 16) Widespread in southern Europe: Spain, south- 208 ern France, Italy, Sardinia, Austria, Czechoslovakia, Hungary, Yugoslavia and Crimea and in western Asia: Anatolia, Armeniya and western Turkmeniya (Kopet-Dag range). Etymology A noun in apposition. From me/anos (Greek), black and ptera (Greek), wings, referring to the black androconial scales on the forewing underside in the males. PHYLOGENETIC AND BIOGEOGRAPHIC CONSIDERATIONS The two species discussed here clearly belong to Trifurcula Zeller s. |., since they possess four out of the six apomorphies for this taxon as listed by van Nieukerken (1986b: 63). Character 44 (velvet patch of raised androconial scales on male hind- wing-underside) is absent in both species, as is also the case in Trifurcula (Glaucolepis) saccharella (Braun, 1912), the type species of Glaucolepis. Since this patch is present in all other Trifurcula species, examined by us, it has most likely been secondarily lost in these three species. The doubtful apomorphy 45 (Hostplant: Fabaceae) has already been discussed by van Nieukerken (1986b). The T. raikhonae species group is here regarded as belonging to Glaucolepis, with which it shows the closest similarity. Van Nieukerken (l.c.) only lists three apomorphies for Glaucolepis: the spe- cies under discussion show some deviations in these characters: 48. Transverse bar of transtilla lost. In both species the transverse bar is apparent, although it is not present as a sclerotized bar. We tentatively assume that this can be regarded as a first step towards the complete loss of the transverse bar. 49. Aedeagus with spines near phallotrema: these are completely lacking in the razkhonae group. 50. Vesica with single long cornutus: the vesica here has indeed one long cornutus, but unlike other Glaucolepis species studied, in T. melanoptera there are some additional small cornuti present. We believe that the condition of character 48 and 50 in the ratkhonae group and the large similarity of the male genitalia in general are sufficient for inclusion in Glaucolepis. On this basis we also have synonymized Sinopticula Yang. However, it is not unlikely that this group belongs to a clade, which is the sistergroup to all remaining known species of Glaucolepis. Character 49 could then be an apo- morphy for the remaining species. An additional apomorphy for Glaucolepis, in- cluding the species under study, is possibly the long, straight basal part of the ductus spermathe- cae, proximal to the coiled part. In Trifurcula s. str. and in Levarchama Beirne, the coiled part starts almost immediately near the vestibulum. The close similarity in many characters between T. raikhonae and melanoptera makes it very likely that both share a common ancestor, which once was widely distributed in the Palaearctic. The spe- cies now have a completely vicariant distribution: melanoptera has a distribution type which very much resembles those of holomediterranean ele- VAN NIEUKERKEN & PUPLESIS: Trifurcula raikhonae group ments of the expansive type (De Lattin 1967), whereas T. raikhonae is widespread in the Central Asian mountains. Although melanoptera occurs as far east as the Kopet-Dag mountains in Turkme- niya, the deserts between this range and the Cen- tral Asian mountains (southern parts of the Kar- akum) form an important gap, which is apparent in the distributions of many taxa (Kryzhanovskij 1965, De Lattin 1967, Matyushkin 1982). Although the present-day desert could be an important bar- rier, the separation most likely goes back to the Pleistocene, when Europe and Central Asia were separated by the enlarged Aralo caspian Sea and the West Siberian Ice lake, or extensions of the continental ice-shields (De Lattin 1967, see also review in Tangelder 1988). ACKNOWLEDGEMENTS We would like to thank the following persons for the loan of specimens and the permission to retain some duplicates: G. Baldizzone (Asti, Italy), R. Buvat (Marseille, France), G. Derra (Bamberg, Germany), C. Gielis (Lexmond, Netherlands), P. Ivinskis (Vilnius, Lithuania), R. Johansson (Växjö, Sweden), O. Karsholt (ZMUC, Copenhagen), the late F. Kasy (NHMw, Vienna, Austria), J. H. Kuch- lein (Wageningen, Netherlands), V. I. Kuznetsov (ZMAS, St. Petersburg, USSR), G. R. Langohr (Sim- pelveld, Netherlands), A. and Z. Lastivka (Brno, Czecho-Slovakia), G. Luquet (MNHN, Paris, France), A. L. Lvov’skii (ZMAs, St. Petersburg, USSR), S. Yu. Sinev (ZMAS, St. Petersburg, USSR) and M. Toth (Plant protection institute, Budapest, Hungary). A. van Assen is acknowledged for assist- ance with operating the SEM, I. Henneke for pho- tographic assistance. R. de Jong and J. van Tol are acknowledged for their comments on the manus- cript. REFERENCES Arnett, R. H. & G. A. Samuelson, 1986. The insect and spider collections of the world. 220 pp. - E. J. Brill/Flora & Fauna publications, Gainesville. Kryzhanovskij, O. L., 1965. Sostav i proiskhozhdeniye nazemnoj fauny Srednej Azii. - Moskva-Leningrad, 381 pp. [The composition and origin of the terrestrial fauna of Central Asia]. Lattin, G. de, 1967. Grundriss der Zoogeographie. — Gus- tav Fischer Verlag, Stuttgart. 602 pp. Matyushkin, E. N., 1982. Regional naya differenciyaciya lesnoi fauni Palearctiki v proshlom i nastoyashchem. - In: Teoreticheskiye i prikladniye aspekti biogeogra- fii. Moskva, Nauka: 52-80. [Regional differentiation of the forest fauna of the Palaearctic in past and present. - Theoretical and applied aspects of biogeo- graphy |. 209 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Nieukerken, E. J. van, 1986a. A provisional phylogenetic check-list of the western palaearctic Nepticulidae, with data on hostplants (Lepidoptera). - Entomolog- ica scandinavica 17: 1-27. Nieukerken, E. J. van, 1986b. Systematics and phylogeny of Holarctic genera of Nepticulidae (Lepidoptera, Heteroneura: Monotrysia). - Zoologische Verhande- lingen, Leiden 236: 1-93. Nieukerken, E. J. van, 1990. The Trifurcula subnitidella group (Lepidoptera: Nepticulidae); taxonomy distri- bution and biology. - Tijdschrift voor Entomologie 133: 205-238. Nieukerken, E. J. van, E. S. Nielsen, R. Johansson & B. Gustafsson, 1990. Introduction to the Nepticulidae. - In: R. Johansson et al., The Nepticulidae and Oposte- gidae (Lepidoptera) of North West Europe. - Fauna Entomologica Scandinavica, 23: 11-109. CORRIGENDA Puplesis, R. K., 1985. Novye vidy molej-maljutok (Lepi- doptera, Nepticulidae) s juga dal’nego vostoka i Tadz- hikistana (New species of the nepticulid moths from Southern far east and Tadzhikistan). - Trudy zooli- cheskogo Instituta, Akademiya Nauk SSSR, Leningrad 134: 59-72. Tangelder, I. R. M., 1988. The biogeography of the Ho- larctic Nephrotoma dorsalis species-group (Diptera, Tipulidae). - Beaufortia 38: 1-35. Yang, Chi-kun, 1989. Sinopticula sinica (Lepidoptera: Nepticulidae), a new genus and species from China. - Entomotaxonomia 11: 79-82 [in Chinese and Eng- lish]. Received: 29 October 1991 Accepted: 4 November 1991 Nieukerken, E. J. van, 1990. The Trifurcula subnitidella group (Lepidoptera: Nepticuli- dae): taxonomy, distribution and biology. - Tijdschrift voor Entomologie 133: 205-238. Unfortunately the following errors have been overlooked during proof-reading of the above cited paper: p. 207. - Figs. 7-10: figs 9 and 10 have been accid- entally interchanged, the left figure with no. 9 actually is fig. 10 (T. coronillae), the right one is fig. 9 (T. subnitidella). Arrows in figs 8-10 have been omitted. p. 222. - Figs. 57-60. Abbreviations: bs=black scales; cf=costal fold; fw=forewing; hw=hind- wing; yp=yellow patch. p. 228. - The sentences after the last paragraph of p. 228 (male genitalia of zberica) were accidentally omitted during page formatting: 210 [sublat-]eral processes. Aedeagus 335-340 um long, with ventral carina fringed; aedeagal tube posteriorly spatulate, dorsal lobe at right side con- spicuous, with serrate margin; vesica with one long spine-like cornutus (125-145 um), with blunt tip, joined basally to a conical cornutus (50 um); further a large cornutus with serrate tip; very few long spine-like cornuti and numerous small ones. Juxta fig. 78. p. 230. - Line 1-2, right column: read hind-wing in stead of hindwing. In some holotype designations, the genitalia slide number has not been mentioned, they are: p. 219, 5. T. victoris: Genitalia slide EVN 2743. p. 225, 7. T. josefklimeschi: Genitalia slide EvN 2744. p. 228, 8. T. zberica: Genitalia slide EvN 1928. p. 230, 9. T. silviae: Genitalia slide EVN 2742. PJOTR OOSTERBROEK & Br. THEOWALD Institute of Taxonomic Zoology, Amsterdam PHYLOGENY OF THE TIPULOIDEA BASED ON CHARACTERS OF LARVAE AND PUPAE (DIPTERA, NEMATOCERA) with an index to the literature except Tipulidae Oosterbroek, P. & Br. Theowald, 1991. Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera), with an index to the literature except Tipulidae. - Tijdschrift voor Entomologie 134: 211-267, figs. 1-180. [1ssN 0040-7496]. Published 18 December 1991. Phylogenetic relationships of the Tipuloidea (the families Limoniidae, Cylindrotomidae and Tipulidae) are discussed on the basis of larval and pupal characters. In comparing outgroups, special attention is given to the synapomorphies listed by Wood & Borkent (1989) for the infraorder Psychodomorpha. In total 105 characters of larvae and pupae are evaluated. A phylogeny is presented for the Tipuloidea as well as for the genera of the subfamilies Eriopterinae, Hexatominae, Pediciinae and Limoniinae. An index to the literature on the immature stages except Tipulidae is added. Correspondence: Dr. P. Oosterbroek, University of Amsterdam, Institute of Taxonomic Zoology, Department of Entomology, Plantage Middenlaan 64, 1018 DH Amsterdam, The Netherlands. Keywords. — Phylogeny, Tipuloidea, Psychodomorpha, Nematocera, larvae, pupae. CONTENTS (lntroductio NEA A eee 211 DapMCthod sents MIT ER RURAL LA be eb Ed 212 SRO UELEOUPSI EN OI Pr A tL Ue 213 CITI pülo idea ra ENI ER I 217 DE TIOpterinaeyys. PET LEE eo dee ee 232 Gmiblexatominae ae: 238 VRR edicinaene een et NOMME D ARE 242 SMlimoniinaer nt een 245 OmRemainingrcenerass nen ee eee 245 1OMIindex tothe literature Nm 252 MÉPACKknowiedsements AREN soc 260 IONRieferences MERE e RAI LES 2 O 260 1. INTRODUCTION The primary concern of this paper are the char- acters of larvae and pupae of Tipuloidea and their contribution to our understanding of phylogenetic relationships among and within the major groups. Translation of the available information into a for- mal classification is considered premature for sev- eral reasons. Characters of the adults are not yet evaluated phylogenetically. Moreover, immature stages are known for representative species of about 150 genera and subgenera, mainly from the Northern Hemisphere, which is about one-third of the recognized genera and subgenera worldwide. Progress in our knowledge of the immature stages of Southern Hemisphere taxa is very important and might well influence certain aspects of the postulated relationships. Tipuloidea as discussed in this paper is identical to: (a) the superfamily Tipuloidea sensu Hennig (1973), comprising the families Limoniidae, Cylin- drotomidae and Tipulidae, (b) the family Tipulidae s.l. of most North American and British workers, who usually recognize one family with three sub- families, (c) the infraorder Tipulomorpha sensu Wood & Borkent (1989). The family Limoniidae is, with about 10,000 described species, one of the largest families of Diptera. Five subfamilies are usually recognized. Immature stages of one subfamily, the Lechriinae (three genera with a limited number of species in Africa, SE Asia and Australia) are not known. The other four subfamilies, Eriopterinae, Hexatomi- nae, Pediciinae and Limoniinae, are discussed 211 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 below. The tribal arrangement of the Palaearctic genera of Limoniidae is given by Dienske (1987), Savchenko (1989) and Savchenko et al. (1992). In Cylindrotomidae about 70 species have been described, in two subfamilies, Cylindrotominae and Stibadocerinae. Information about the immature stages of the latter subfamily (four small genera in SE Asia and Chile) is not available. The family Tipulidae accounts for some 4100 described species. Holarctic genera are assigned by some workers to three subfamilies, Dolichopezi- nae, Ctenophorinae and Tipulinae. This subdivi- sion is not corroborated by the phylogenetic rela- tionships and will be discussed in a forthcoming paper. The superfamily Tipuloidea is without doubt monophyletic (Hennig 1973, Wood & Borkent 1989), among others on the basis of larval charac- ters. A phylogenetic scheme for the Tipuloidea was first framed by Alexander (1920), without a clear justification but apparently based on information from all life-stages. A different phylogeny was presented by the same author in 1927, based on the radial field of the wing. This study was criticized by Hennig (1954), who did not agree with some of Alexander's interpretations. A phylogeny based on pre-imaginal and imaginal characters of Palaearc- tic taxa was presented by Savchenko (1966, see also Savchenko 1979, 1983b), who concluded that, al- though Limoniidae and Cylindrotomidae are ap- parently sistergroups, there also exists a close re- lationship between Tipulidae and Limoniidae. Both Alexander and Savchenko started from the concept that the three families within the Tipuloi- dea are monophyletic. This view was not followed by Hennig, especially not with respect to the Limo- niidae, and he suggested that the Tipulidae might well be the sistergroup of part of the Limoniidae. Oosterbroek (1986) proposed a new classifica- tion of the Tipuloidea into four families based on a preliminary study of larval and pupal characters. These four families and their sistergroup relations are: Cylindrotomidae - remaining Tipuloidea; Li- moniidae - Eriopteridae + Tipulidae; Eriopteridae - Tipulidae. This classification is not confirmed by the results of the present study. 2. METHODS Information about larval and pupal characters was available for those species of Limoniidae and Cylindrotominae mentioned in the index. Most characters are taken from the literature. Only a limited amount of characters could be studied from preserved material (see index). Valuable informa- tion about the morphology of larvae and pupae in Diptera, Nematocera and Tipuloidea is presented 212 by Alexander 1920 (Tipuloidea), Alexander & By- ers 1981 (Tipuloidea), Anthon 1943a (Nematoc- era), Bangerter 1928-1934 (Tipuloidea), Bengts- son 1897 (Cylindrotominae), Brauns 1954a, 1954b (Diptera), Brindle 1960 (Tipulidae), 1967 (Limo- niidae and Cylindrotominae), Byers 1961 (Dolichopeza), 1983 (Chionea), Chiswell 1955 (Ti- pulidae), Cook 1949 (Diptera), Courtney 1990, 1991 (Blephariceromorpha), Cramer 1968 (Tipu- loidea), Crisp & Lloyd 1954 (Nematocera), Gelhaus 1986 (Tipulidae), Gérard 1968 (Pilaria), Giljarov (ed.) 1964 (Insects), Griffiths 1990 (Nematocera), Hennig 1948-1952, 1973 (Diptera), Hinton 1954- 1968 (respiratory horns), Hynes 1958-1990 (Limo- niidae), Krivosheina 1964 (Tipuloidea), 1969 (Diptera), Lenz 1920a (Thaumastoptera), Lindner 1959 (Limoniidae), McAlpine (ed.) 1981, 1989 (Diptera), de Meijere 1917 (Diptera), Oldham 1926 (Pedicia), Peterson 1960, 1965 (Insects), Peus 1952 (Cylindrotominae), Pokorny 1978 (Eloeophila), Reusch 1988 (Limoniidae), Rogers 1926-1933 (Tipuloidea), Rozkosny 1980 (Tipuli- dae, Cylindrotominae), Rozkosny & Pokorny 1980 (Limoniidae), Smith 1989 (Diptera), Teskey 1981 (Diptera), Theowald 1957, 1967 (Tipulidae), To- kunaga 1930, 1933 (Dicranomyia), Vaillant 1951 (Orimarga), 1953 (Symplecta), Wiedenska 1987 (Phylidorea), Wirth & Stone 1956 (Diptera), Wolff 1922 (statocysts), Wood & Borkent 1989 (Nemat- ocera), Wood 1952 (Tipuloidea). Relationships are determined according to the phylogenetic principles as worked out by Hennig (1966), and as modified by Wiley (1981). Decisions about character polarities are based, as far as pos- sible, on outgroup rules (Watrous & Wheeler 1981, Maddison et al. 1984). The terminology largely follows Teskey (1981). The larval characters discussed and figured are those of the full grown, last larval stage, unless stated otherwise. POLYNEURA OLIGONEURA Mr (REMAINING È DIPTERA) n a AURA o (>) = 2e =) (©) a 2 F Fr Fig. 1. Phylogeny of Tipuloidea, Trichoceridae and the remaining Diptera. For explanation see text. 3. OUTGROUPS With respect to outgroup comparison, several levels can be recognized. The first level deals with the immediate sistergroup relation between Tipu- loidea and other Nematocera, the second with the position of this sistergroup among the Nematoc- era, the third with the sistergroup of the Nemat- ocera. First level outgroups In the literature the sistergroup of the Tipuloi- dea are either the Trichoceridae (winter crane flies; Hennig1968, 1973; Griffiths 1990) or all other Diptera (Wood & Borkent 1989). Tipuloidea and Trichoceridae as sistergroups (fig. 1) Hennig (1968, 1973) united Tipuloidea and Tri- choceridae in a monophyletic group (his suborder Polyneura or infraorder Tipulomorpha) on the ba- sis of a number of synapomorphies. Wood & Bor- kent (1989) presented a critical analysis of these synapomorphies, and concluded that some are to be rejected or of little phylogenetic importance, while others are doubtfully homologous or in need of further investigation. Arguments in favour of the monophyly of the Polyneura have been put forward by other authors as well (Rohdendorf 1964, Dahl & Alexander 1976, Dahl 1980, Griffiths 1990). The synapomorphies mentioned by Griffiths (1990) are (fig. 1): (1.) Reduction of male cerci, (2.) Development of go- nopods from posterolateral zones of proliferation, (3.) Female cerci with single article, (4.) Only three branches of radial sector reaching wing margin, (5.) Forward displacement of distal section of M2. Tipuloidea and ‘all other Nematocera’ as sister- groups (fig. 2) In Nannochoristidae, one of the important out- groups of the Diptera (see fig. 3), the larval mand- ible is provided with a separate sclerotized lobe (the lacinia mobilis of Pilgrim 1972). Among Ti- puloidea a positionally similar structure is found only in Cylindrotominae (fig. 162), Tipulidae (figs. 179, 180) and the hexatomine genera Pilaria and Ulomorpha (fig. 61), the only two genera of Tipu- loidea with a twofold mandible. In their comprehensive analysis of the phyloge- netic relationships among the Nematocera, Wood & Borkent (1989) tentatively considered the pres- ence of a ‘lacinia mobilis’ in Nannochoristidae and OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Tipuloidea a symplesiomorphy and its absence in all other Nematocera a synapomorphy (fig. 2: char- acter 1). However, the lacinia mobilis of Nanno- choristidae and the separate sclerotized lobe of Cylindrotominae and Tipulidae most probably are not homologous. In Nannochoristidae the lacinia mobilis is already present in the first instar larva and the prosthecal brush of setae is situated basally (Pilgrim 1972: fig. 6). The separate sclerotized lobe of Cylindrotominae and Tipulidae is not yet pres- ent in the first instar larvae (fig. 161) and the prosthecal brush is situated at the apex (figs. 162, 179, 180). The only remaining account of a ‘lacinia mobilis’ in Tipuloidea is the separated sclerotized lobe in Pilaria and Ulomorpha (fig. 61: pr). This lobe is not toothed as in Nannochoristidae. In the first stage larva it is already present, whereas the mand- ibles are not yet bifold (Gérard 1968 for Pilaria). The phylogenetic position of these two genera (fig. 6) and the very unique, highly specialized structure of the mandibles, which can close on itself like a ‘pocket knife’ (Crisp & Lloyd 1954), indicate that presence of a separated sclerotized lobe in Pilaria and Ulomorpha is most probably a synapomorphy. The same might be true for the lacinia mobilis of the Nannochoristidae because it is not present in the other Mecoptera. Courtney (1990, 1991) assumed that the separ- ate sclerotized lobe is homologous with the subba- sal, thumb-like projection found in Psychodomor- pha (see below, character 42). This subbasal projection, present in a number of Tipuloidea, oc- curs in Cylindrotominae together with the separate sclerotized lobe (fig. 162). Therefore, the pre- sumed homology might be correct for other Nematocera, but can not be maintained for the Cylindrotominae and Tipulidae. In his review of the paper by Wood & Borkent (1989), Griffiths (1990) hesitated to accept the absence of a separate sclerotized lobe as a synapo- morphy for all Nematocera except Tipuloidea. In the present study, the separate sclerotized lobe is interpreted as a synapomorphy for Cylindrotomi- nae and Tipulidae (chapter 4: character 22). The phylogenetic position of these taxa (fig. 4) does not favour the assumption that this character belongs to the groundplan of the Tipuloidea. It can be concluded that character 1 of figure 2 is a questionable synapomorphy for all Nemato- cera except Tipuloidea. During the present study, a sistergroup relation between the Tipuloidea and all other Nematocera could not be established on the basis of other synapomorphies. 213 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 PSYCHODOMORPHA ? - +-+--- + + + +++ - APHOHEHHEPAAPAPPAMA N =x fa) @ (e) O) DEUTEROPHLEBIIDAE BLEPHARICERIDAE AXYMYIIDAE PERISSOMMATIDAE ANISOPODIDAE PTYCHOPTERIDAE MYCETOPHILIDAE DIXIDAE NYMPHOMYIIDAE PACHYNEURIDAE BIBIONIDAE SCIARIDAE CECIDOMYIIDAE PSYCHODIDAE TRICHOCERIDAE SCATOPSIDAE SYNNEURIDAE TANYDERIDAE TIPULIDAE CORETHRELLIDAE CHAOBORIDAE CULICIDAE Fig. 2. Phylogeny of the Nemat- ocera after Wood & Borkent (1989). Row 1, referring to the pu- pal leg sheaths and row 2, referring to the respiratory system, are added. - Row 1: - pupal leg sheaths side by side, + pupal leg sheaths superimposed, *at least two pairs of leg sheaths side by side (Pty- chopteridae). - Row 2: M meta- pneustic, a apneustic, A amphi- pneustic, P peripneustic, HO holopneustic, HE hemipneustic, * except Archaeochlus (Chirono- midae). THAUMALEIDAE SIMULIIDAE CERATOPOGONIDAE CHIRONOMIDAE Second level outgroups Oligoneura sensu Hennig (1968, 1973) (fig. 1) Hennig (1968, 1973), following Lindner (1949), divided the Diptera in two suborders, Polyneura (Tipuloidea plus Trichoceridae) and Oligoneura (the remaining Diptera). Oligoneura can be consi- dered monophyletic on the basis of the following characters (fig. 1). (6.) Arolium absent, replaced by a flap-like empodium, (7.) A2 rudimentary or ab- sent, (8.) Base of veins M and R connected by an oblique crossvein. Characters 6-8 were discussed by Wood & Bor- kent (1989), who concluded that all three charac- ters are in need of further investigation and that other evidence to corroborate the monophyly of the Oligoneura seems to be lacking. After a detailed study of the fifth tarsomere throughout the Dip- tera and a number of Mecoptera, Röder (1984) concluded that character 6 is a synapomorphy for the Oligoneura. Larval mouthparts in Tipuloidea do not show the striking similarities as present between Tri- choceridae, Psychodidae, Anisopodidae and Sca- topsidae, discussed in more detail in the next chap- ter. These similarities must be interpreted as belonging to the groundplan condition of the Nematocera, if the phylogeny presented in fig. 1 is correct. A similar conclusion was reached by Ed- wards (1926, 1928), Anthon (1943a), Hennig 214 (1954), Dahl & Alexander (1976). Edwards (1928) was unable to discover anything among the adults which would suggest that Trichocera should be included in the Anisopodidae (‘In almost every respect the adults of the genus [ Trichocera] are typical Tipulids ... It seems probable that we should regard Trichocera as a primitive genus which in its larvae at least has retained many of the characters of the ancestors of the Tipulidae, and its resemb- lances to the Anisopodidae are probably due to this fact’). Some of the characters of the larval mouthparts are also present in Ptychopteridae and Tanyderi- dae. Therefore, on the basis of fig. 1, these families together with Psychodidae, Anisopodidae and Sca- topsidae are to be regarded as second level out- groups. Psychodomorpha sensu Wood & Borkent (1989) (fig. 2) Keilin (1912) was the first to decide that on larval characters Trichocera should be separated from the Tipulidae s.l. Wood & Borkent (1989) analysed these larval characters in much detail, with emphasis on the larval mouthparts. Several of these mouthpart characters were interpreted by Wood & Borkent as synapomorphies (fig. 2: char- acters 38-45), uniting Trichoceridae, Psychodidae, Anisopodidae and Scatopsidae in the infraorder Psychodomorpha, together with Perissommatidae and Synneuridae. Griffiths (1990) stated that the synapomorphies between Tipuloidea and Trichoceridae (fig. 1: char- acters 1-5) can not easily be dismissed. Therefore, in his view, the Tipuloidea are to be included in the Psychodomorpha as sistergroup of the Trichoceri- dae, stating that this: ‘is reconcilable with Wood & Borkent’s justified emphasis on the synapomor- phies between the larval head structure of Tri- choceridae and that of other Psychodomorpha, if we assume that the considerable diversity of head structures shown by the larvae of Tipulidae s.l. represents a transformation series from a ground- plan structure similar to that of Trichocera’. According to the above, the other families in- cluded in the Psychodomorpha, especially Psycho- didae, Anisopodidae and Scatopsidae, are to be con- sidered second level outgroups. Synapomorphies for the Psychodomorpha The conclusion about second level outgroups is more or less the same for both phylogenies pres- ented in fig. 1 and fig. 2. In both instances, charac- ters of the larvae of Tipuloidea must be interpreted as representing a transformation from a ground- plan structure similar to that of Trichoceridae, Psy- chodidae, Anisopodidae and Scatopsidae. Whether this interpretation is correct can be determined by comparing the larval characters of Tipuloidea with the synapomorphies listed by Wood & Borkent (1989) for Psychodomorpha, Ptychopteromorpha and Culicomorpha (fig. 2: character 5) and Psycho- domorpha (fig 2: characters 38-45): Character 5 (Wood & Borkent): ‘Epicondyle dis- placed medially and hypocondyle displaced later- ally, with the mandibles operating obliquely or vertically’. — As discussed below (character 41), this synapomorphy is probably of little phylogenetic significance. Courtney (1990, 1991) accepted it with some reservation to support the monophyly of Blephariceromorpha and Psychodomorpha. Character 38: ‘Labrum conical or wedge-shaped, rounded, or pointed anteroventrally, more or less covered apically and ventrally with backwardly projecting hairs’. - In many Tipuloidea the labrum is broadly rounded and especially ventrally covered with backwardly projecting hairs. A distinctly nar- row labrum is found in Eriopterinae (figs. 13, 19) and Cylindrotominae. The plesiomorphic condi- tion, labrum flattened dorsoventrally and usually bilobate’ is present in most Hexatominae, Limonii- nae and Tipulidae (e. g. fig. 73). The apomorphic condition is present also in Blephariceridae, Axy- myiidae and Thaumaleidae. In Psychodomorpha some Psychodidae (Phlebotomus, Trichomyia) and the Synneuridae exhibit the plesiomorphic condi- tion. OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Character 39: ‘Premandible in the form of a rounded or oval sclerite with a comb-like row of evenly spaced, anteriorly projecting teeth, presum- ably operated by the labral retractor muscles to comb the mandible or maxilla’. - In Tipuloidea premandibles as in Psychodomorpha are absent. Especially in Hexatominae, however, the epipha- rynx is provided with two relatively large sclero- tized plates situated caudal of the epipharyngeal bar (fig. 73; see also Pokorny 1978: figs. 14-16 for Eloeophila and Wiedenska 1987: figs. 4-5 for Phy- lidorea). Positionally these plates might be homol- ogous to the premandibles. The same applies for the structures figured by Vaillant (1953: fig. 4) for the eriopterine genus Symplecta. Among Psycho- domorpha, the apomorphic condition is not pres- ent in the anisopodid genus O/biogaster (Keilin & Tate 1940, Anthon 1943b), the psychodid genus Trichomyia and the Synneuridae (Wood & Borkent 1989). Wood & Borkent (1989) considered the shape of the premandibles as ‘one of the most convincing pieces of evidence of the monophyly of the infraorder [Psychodomorpha]’, although they did not indicate which taxa exhibit the plesiomor- phic condition (‘premandibles in the form of a simple, unadorned sclerite, serving only as a point of insertion of the labral retractor muscles’). Ble- phariceromorpha, Axymyiidae and Bibionomor- pha apparently lack premandibles except for Myce- tophilidae (s.1.) and Sciaridae. Premandibles in the two last-mentioned families are not necessarily ho- mologous with those present in Psychodomorpha (Goetghebuer 1925, Edwards 1926, Plachter 1979). Edwards (1926) and Courtney (1990) supposed that premandibles possibly arose as a groundplan feature of Diptera larvae. This might be the correct interpretation because this character should be placed at a lower level in the phylogeny of figure 2, namely at the same position as character 5 (see also Courtney 1991: fig. 4b). This lower position is apparent from the statement by Wood & Borkent (1989) that the premandible of the Psychodomor- pha represents the plesiomorphic condition with respect to the apomorphic premandibles of Pty- chopteromorpha and Culicomorpha (fig. 2: charac- ter): Character 40: ‘Torma articulated with the dorsal labral sclerite’. - This character is present in sev- eral groups of Tipuloidea, most distinctly in the larger Tipulidae (fig. 174), but observed also in Dactylolabis and several Limoniinae. Articulating tormae are mentioned in the literature for Crypto- labis (Eriopterinae, Hynes 1963), Pedicia (Pedi- ciini, Wardle & Taylor 1926), Dicranomyia (Limo- niinae, Tokunaga 1933), and similar structures are figured and described for Thaumastoptera (Limo- niinae, Lenz 1920a). Cook (1949) identified the 215 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 premandibles in Holorusia, but given the position of these sclerites it is likely that they are the tormae (fig. 174). According to Wood & Borkent (1989) this character might represent a synapomorphy at the same position in figure 2 as character 5, whe- reas Courtney (1990, 1991) unites Blepharicero- morpha and Psychodomorpha on the basis of the presence of articulated tormae in Blepharicero- morpha. Character 41: Mandibular movement ‘in a nearly vertical plane as a result of a shift in position of the condyles, with the mandibles striking the anterior edge of the hypostoma or hypostomal teeth’. - An oblique or vertical orientation of the larval mand- ibles is present in all Eriopterinae, Hexatominae, Ula and sometimes Pediciini (chapter 4: character 10). The mandibles move in a horizontal or nearly horizontal plane in all other Tipuloidea, except Cylindrotominae. In this subfamily the mandibles move horizontal in first stage larvae and almost vertical in second stage to full grown larvae (Bengtsson 1897, Alexander 1920, Peus 1952). Among Nematocera as a whole this character is subject to homoplasy and probably of questionable value in phylogenetic interpretations (Courtney 1990, 1991). As stated by Wood & Borkent (1989: character 5), the positions of the mandibular artic- ulation points have probably shifted more than once and reversals have undoubtedly occurred. The orientations of the mandibles vary in the Psycho- domorpha (horizontal in the psychodid subfami- lies Trichomyiinae (Wood & Borkent 1989) and Bruchomyiinae (Satchell 1953), obliquely in Peris- sommatidae), Ptychopteromorpha (Tanyderidae horizontal, some Ptychopteridae vertical), and Cu- licomorpha (horizontal in Stenochironomus). In most Bibionomorpha the mandibles move in a ho- rizontal plane but in Mycetophilidae (s.l.) the mandibles operate in a semi-circular way (Plachter 1979: "halbkreisförmig'). Character 42: ‘Mandible in the shape of a chela with a small, subbasal, thumb-like projection curv- ing toward the apex of the mandible’ (figs. 9, 10). — Mandibular shapes are rather varied within Ti- puloidea. A small subbasal projection curving to- ward the apex is present in a few Eriopterinae (fig. 11; see also chapter 5: character 56) and in some species of Dactylolabis (fig. 128). Within Psycho- domorpha the apomorphic condition is not found in the psychodid genera Bruchomyia and Phlebo- tomus (Satchell 1953), and is less distinct in Per- issommatidae (Wood & Borkent 1989) and the scatopsid subfamily Ectaetiinae (Laurence 1953). Character 43: ‘Apical portion of mandible separ- ated from the base by a less strongly sclerotized line of weakness’. - Bifold mandibles of a very charac- teristic and completely different shape than in Psy- 216 chodomorpha (figs. 9, 10) are present in the highly specialized hexatomine genera Pilaria and Ulo- morpha (fig. 61). In all other Tipuloidea the apical portion is solidly fused with the base of the mand- ible (e. g. fig. 11). According to Anthon (1943a) and Hennig (1973) bifold mandibles probably be- long to the groundplan condition of the Diptera and after Lindner (1959) its occurrence in Tipuloi- dea must be interpreted as a reversal to the plesi- omorphic condition. If apomorphic, this character unites Psychodomorpha and Blephariceromorpha (Courtney 1990, 1991) and possibly also Ptychop- teromorpha (Tanyderidae: Anthon 1988: M:- schoderus; Ptychopteridae: Anthon 1943a, Wood & Borkent 1989). A bifold mandible is also present in one subfamily of Psychodidae (in Psychodinae, where it is similar to the other Psychodomorpha). In the other three subfamilies of Psychodidae (Bru- chomyiinae, Phlebotominae, and Trichomyiinae) the mandible is simple (Satchell 1953, Quate & Vockeroth 1981). The apomorphic condition could not be ascertained for the Perissommatidae and is apparently absent in the Synneuridae (Wood & Borkent 1989) and the scatopsid subfamily Ectae- tiinae (Laurence 1953). Character 44: ‘Cardo reduced, mainly membran- ous’. - The description of this synapomorphy by Wood & Borkent (1989) does probably not refer to the cardo itself, which is a well developed sclerite in several Anisopodidae, Trichoceridae and Scatop- sidae (see Anthon 1943a: figs. 55-60, Krivosheina 1969), but to the pattern of ‘backwardly pointing setae forming a characteristic pattern, unique in the Diptera’, situated on the membranous area between the cardo and the maxillar lobes. These backwardly pointed setae are absent in Tipuloidea, but also in the anisopodid genera Mycetobia and Olbiogaster (Keilin 1919, Anthon 1943a, 1943b, Keilin & Tate 1940, Krivosheina 1969), whereas similar setae are present in Philosepedon (Psycho- didae) and Ptychoptera (Ptychopteridae) (Anthon 1943a). A membranous cardo occurs in Nannocho- ristidae and Blephariceromorpha and is considered further evidence for the sistergroup relation be- tween Blephariceromorpha and Psychodomorpha by Courtney (1990, 1991). In Tipuloidea the cardo is usually well developed and of the same shape as in many other Nematocera (see also Cook 1949). Little information is available for Eriopterinae and Hexatominae, but in Cryptolabis (Hynes 1963) and Pseudolimnophila (fig. 58) the cardo lies parallel to the ventral anterior margin of the genae, as in the first and second level outgroups. In Ula and the members of the Dactylolabis-Tipulidae lineage of figure 4, only the proximal end of the cardo is near to the anterior margin of the genae (e. g. figs. 160, 173). This character, which can not be interpreted MECOPTERA NANNOCHORISTIDAE SIPHANOPTERA DIPTERA MECOPTERA NANNOCHORISTIDAE SIPHANOPTERA DIPTERA MECOPTERA NANNOCHORISTIDAE for the reduced cardo of Pediciini (fig. 100), seems to be closely associated with the orientation of the anterior margins of the genae (chapter 4: character DI Character 45: ‘Maxillary palpus reduced to a disc- like sclerotized ring’. - According to Wood & Bor- kent (1989), in the Trichoceridae, Perissommati- dae, Scatopsidae, and Anisopodidae, the maxillary palpus is flush with the surface of the maxilla, recognizable only by a ring of chitin surrounding the sensilla. This type of maxillary palpus is found in Eriopterinae (figs. 15, 23), Hexatominae, Pedi- ciini (the two last-mentioned taxa with the outer maxillary lobe elongated and the sensilla situated near or at the apex, figs. 68, 100), Ula (fig. 113) and Dactylolabis. In other Tipuloidea the sclerotized ring surrounding the sensilla is cylindrical and us- ually well developed (fig. 121; see also chapter 4: character 12). A reduced maxillary palpus occurs also in Cecidomyiidae (Mamaev & Krivosheina 1965), Mycetophilidae (s.1.) and Sciaridae (Plachter 1979). Conclusions The above comparison shows that a number of larval head characters of Tipuloidea, especially in Eriopterinae and Hexatominae, correspond to the synapomorphies listed by Wood & Borkent (1989) for the Psychodomorpha. From this it can be con- cluded that Edwards (1926, 1928) and Griffiths (1990) were probably right when they stated that larval characters of Tipuloidea are to be interpreted as a transformation series from a groundplan struc- ture similar to that of Trichocera. It is also apparent that several of the synapom- orphies listed by Wood & Borkent (1989) for the Psychodomorpha are not present in all represen- tatives of this infra order and/or are present in other families of Nematocera as well. Some of these characters, therefore, must be regarded OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Fig. 3. Phylogenetic arrangements for the Antliophora. - A, after Wood & Borkent (1989); B and C, after Willmann (1989) and Griffiths (1990). SIPHANOPTERA DIPTERA groundplan structures for a grouping of nematoce- ran families larger than the Psychodomorpha (as worked out for the Blephariceromorpha by Court- ney 1990, 1991), or for even all Nematocera. From this it follows that some of the Psychodomorpha families might belong to the most basal lineages of the Diptera. This latter observation is in agree- ment with the hypothesis by Anthon (1943a) that the presence of a transverse tentorium represents the primitive condition, as found in primitive in- sects, Mecoptera (incl. Nannochoristidae), Sipho- naptera, and among Psychodomorpha in Anisopo- didae and Perissommatidae. Wood & Borkent (1989: 1356), stated that it is difficult to refute the hypothesis that the anisopodid larva represents one of the most primitive types of dipterous head, but on the other hand prefer to suppose that the transverse tentoria of Anisopodidae and Perissom- matidae are a de novo development, apparently not present in the other families of Diptera (see also chapter 4: character E). Third level outgroups (fig. 3) Diptera, Mecoptera (including Nannochoristi- dae) and Siphonaptera are usually united in the Antliophora. Various sistergroup relations have been proposed (fig. 3) but the matter is still open to debate (reviews in Hennig 1981, Willmann 1989, Wood & Borkent 1989 and Griffiths 1990). For the present study, outgroup comparison is lar- gely based on characters of Mecoptera including Nannochoristidae (Byers 1963, Kaltenbach 1978, Pilgrim 1972, Steiner 1930). 4. TIPULOIDEA Introduction On the basis of the available information the Tipuloidea can be divided in at least two, appar- 217 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 < — PEDICI- RN = = Spo NE he SS Een EE nd MES = x = cali Si = (i) a ~ La ze te eo a © 58 Si Ou = ie ee re io A= ie eee aed NT ET SRO oO < = a X = n = x a < (©) D = O =F a. X Lu Lu = A I (o = LU = Lu zE a >) a < Lul a 25 = ST | C] #2 C] [| D 340] Bo [va] L | a 4 | | 5555 [na] E D m B Gi C] m 320 | [| C] [rai 310 C] i [Bi [| L | Sole] Û i Dd C] C] | 290 Û C] a [| L | [| [| mn 28 LJ [ [| [1 270 Bo 26 MN PER Kad had VIN Keri Kad Kad 16 15 13 14 = 12 e 11 7 3 6 10 2. SE —— 1 4E A-G Br ee Fig. 4. Phylogeny of the major = = groups of Tipuloidea. = = os str. ta ei zu RORE MEIER = a =i ui (©) > @ El ol? [a oO [| @ O C] Ein JE ET ET EE L gu 2 8 23 22 21 9 20 19 18 TZ [M CHARACTER NOT PRESENT IN ALL REPRESENTATIVES OF THE TAXON RK] AUTAPOMORPHOUS CHARACTER ently monophyletic groups (fig. 4), the Eriopteri- nae-Hexatominae lineage and the Dactylolabis-Ti- pulidae lineage. These two lineages and the Pedi- ciinae are placed at a basal trichotomy. Characters 4-5 point in the direction of a sistergroup relation between the Pediciinae and the Dactylolabis-Tipu- lidae lineage, character 24 to such a relation be- tween the Pediciinae and the Eriopterinae-Hexato- minae lineage. The monophyly of the Pediciinae, based on characters 6-9, is not corroborated by characters 25-27. Pediciinae are the only Tipuloidea with hairy eyes in the adults, without doubt a symplesiomor- phic character. Therefore, it might be expected to find the Pediciinae at the most basal position, as sistergroup of the Eriopterinae-Hexatominae plus Dactylolabis-Tipulidae lineages. An indication for resolving the trichotomy in this way is the shape of the prementum in Pediciinae (see character 6). It also appears that the pharynx and oesophagus of 218 Pediciini and Ula (detailed description in Miall 1893, Oldham 1926, Lindner 1959) show a certain resemblance to the pharyngeal filter apparatus of Trichoceridae and several second level outgroups (detailed description in Anthon 1943a). Very little information is available for the pharynx and oeso- phagus of other Tipuloidea but a pharyngeal filter apparatus as in Ula (Lindner 1959) and perhaps also Pediciini is not present in Chionea (Eriopte- rinae, Byers 1983), Pilaria and Hexatoma (Hexa- tominae, Cook 1949, Gérard 1968), Antocha and Limonia (Limoniinae, Sinclair, pers. comm.) and Holorusia (Tipulidae, Cook 1949). In the text below, Pediciinae, Cylindrotominae and Tipulidae (s. str.) have the same meaning as the corresponding but sometimes differently ranked taxa in the current systematic literature (Hutson 1980, Dienske 1987, Oosterbroek 1989, Savchenko 1989, Savchenko et al. 1992). On the basis of larval and pupal characters, Eriopterinae O Bee i ae m 9 - = 2 = < ac > © O ee un Se We SEE PEN EREN MENE OE SE SE ER (aye Aelen lesen ray, (Sw) SS U Ze ERN RE < 0 1918 + 5 9 = a 5 ae! PL DE SR mem TE m 82M |] DIVO ara), i] See ONE ISO area 80 Eee Be] (i EE Ss À Ce] M Le ee)" |B): a) |e El LÌ 78 TEU 76 75 74 173 70 VA 69 al 68 67 66 65 64 CSSS 263 Fig. 6. Phylogeny of the Hexatominae. tional and most distinct shortly after ecdysis (Alex- ander 1920, Lindner 1959). The larval respiratory system of Mecoptera is peripneustic, in Nannochoristidae apneustic (first three larval stages) or hemipneustic (full grown larva). Nematocera are usually peri-, amphi- or apneustic (fig. 2), the only other metapneustic forms are found in Ptychopteridae and Culicidae. According to Keilin (1944), first stage larvae of apparently all Diptera are metapneustic (‘The me- tapneustic system appears as the first stage of de- velopment in all respiratory systems except the apneustic one’ (Keilin 1944: p. 31). Preservation of the metapneustic condition in full grown larvae, as in Tipuloidea, therefore, could represent the plesi- omorphic condition. On the other hand ‘metap- neustic and amphipneustic forms which are adapted to a partially submerged condition of life are more specialized and are derived from the primitive terrestrial polypneustic form’ (Keilin 1944: p. 31; see also Hinton 1947: p. 466, Wood & Borkent 1989: p. 1342). B. Larvae hemicephalic Larvae of Tipuloidea are hemicephalic, the head 221 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 capsule can be retracted into the body. This char- acter is also present in the Brachycera. The plesi- omorphic condition in Diptera is without doubt an eucephalic head capsule, as in almost all other Nematocera and related orders. In the nematoce- ran family Axymyiidae the head can be partly re- tracted into the body as well (Mamaev & Krivo- sheina 1966). According to Teskey (1981), the hemicephalic condition in Tipuloidea does not im- ply a vertical orientation of the mandibles as in the Brachycera. It is assumed here that within the Ti- puloidea a horizontal orientation is derived from an oblique orientation (see character 10). C. Ventral margins of the genae extending far an- teriorly Correlated with the hemicephalic condition is reduction of the larval head capsule, whereby in all Tipuloidea the ventral margins of the genae are letting free a large portion of the ventral floor of the head (ventral part of occipital foramen dis- tinctly extended anteriorly). In the related orders and almost all other Nematocera the ventral floor is largely closed or the ventral contact of the genae is situated not further anteriorly than about half- way the length of the genae, some Mycetophilidae (s.1.) excepted. D. Genae posteriorly incised Reduction of the larval head capsule is also ex- emplified by distinct incisions which divide the genae in externo- and interno-lateralia (fig. 14: elp and ilp). These incisions are not present in the related orders or in other Nematocera except for Axymyiidae (Mamaev & Krivosheina 1966). It is noteworthy that in Trichocera the posterior mar- gin of the genae is sharply invaginated at the same position as the incisions of Tipuloidea (see Anthon 1943a: ‘mf’ in fig. 6 & 8). In some species of Ula and in Pediciini the incisions are hardly developed (figs. 93, 110). E. Tentorium reduced A complete tentorium, consisting of tentorial arms and a transverse tentorium is considered to belong to the groundplan of the Diptera by among others Anthon (1943a), Hennig (1973), Teskey (1981). Tentorial arms are present in Trichocera and in several but not all families of Nematocera and Brachycera. In addition to the tentorial arms a transverse connection is apparently restricted to Anisopodidae and Perissommatidae (Wood & Bor- kent 1989), although it is described as well for the nematoceran families Bibionidae (Perraudin 1961), Mycetophilidae (s.l.) (Plachter 1979) and Dixidae (Nowell 1951). In Tipuloidea the trans- verse connection is absent and the tentorial arms 222. TRICYPHONA 2 RHAPHIDOLABINA RHAPHIDOLABIS PARADICRANOTA < = oO x= a > (©) [24 E PEDICIA DICRANOTA Fig. 7. Phylogeny of the Pediciinae. are extremely reduced (Lindner 1959: fig. 9). F. Pupal legs sheaths side by side In all Tipuloidea the pupal leg sheaths are lying side by side. This character must be considered a weak argument for the monophyly of the Tipuloi- dea because several other nematoceran families have the leg sheaths side by side as well. Superim- posed leg sheaths are nevertheless found in all families of the Psychodomorpha (sensu Wood & Borkent 1989) and also in Axymyiidae and Bibio- nidae (fig. 2). G. Hypostoma anteriorly toothed and consisting of central and lateral parts The hypostomal region of Tipuloidea is rather diverse. All taxa, except higher Hexatominae, the majority of Eriopterinae and some species of Ele- phantomyia (see chapter 9), possess a distinctly toothed hypostoma (e. g. fig. 131). Most lower Eriopterinae also possess a toothed hypostoma, although less pronounced (fig. 26; see character 59). Eriopterinae (figs. 12, 13, 15, 16, 26, 33), Hex- atominae (figs. 58, 59) and Pediciini (figs. 94, 97) have the hypostoma divided, in Pediciini with a very characteristic push-button connection (fig. 97), in a few Eriopterinae with a hyaline connec- tion between the anterior ends (Rhabdomastix afra (Wood 1952), Grophomyia jacobsoni (fig. 16, Rog- ers 1927b) and probably also Ewgnophomyia luc- tuosa (Rogers 1928)). In the lower hexatomine genera Pseudolimnophila (fig. 58) and Paradel- phomyia (fig. 59), two, large, separated hypos- tomal plates are present. These are the only two genera of Hexatominae with a more or less com- pact head capsule. The other Hexatominae have the head capsule and the hypostoma reduced further with the ventral rods of the head capsule (which are homologous to the ventral inner mar- gins of the genae) more widely separated (figs. 67, 68). In Ula and the genera of the Dactylolabis-Tipu- lidae lineage the hypostoma is complete, although it is usually only narrowly continuous anteriorly and has a distinct posterior cleft (e. g. fig. 124). In Pseudolimnophila, Paradelphomyia, Ula and a number of genera of the Dactylolabis-Tipulidae lineage the hypostoma consists of two parts, namely a central part situated more ventrally and a lateral part situated more dorsally (figs. 59, 124, 131). The central part is apparently of labial and the lateral part of genal origin (see character 23 for details about the origin of the hypostoma). Outgroup comparison reveals that the hypos- toma consisting of a central and lateral part most probably represents a synapomorphy for Tipuloi- dea. Third level outgroups do not possess a hypos- toma. In Trichocera and some second level out- groups (Anisopodidae: Sylwicola, Mycetobia) a hypostoma is not present but the ventral margins of the genae are connected by a posteriorly situated hypostomal bridge. Other second level outgroups (Psychodidae, Perissommatidae, Anisopodidae: Olbiogaster) as well as a few other families of Nematocera (Ptychopteridae, Culicidae, Simulii- dae, Chironomidae) do possess a complete hypos- toma. In these outgroups the hypostoma appar- ently does not show a subdivision in central and lateral parts (Anthon 1943a, 1943b, Harbach & Knight 1980, Teskey 1981, Peterson 1981). The posterior cleft is very rare among other Nematoc- era and might as well be considered a synapo- morphy (a somewhat similar situation is found in the psychodid species Philosepedon humeralis (Meigen), Anthon 1943a: fig. 17, and the simuliid species Metacnephia saileri (Stone), Peterson 1981: fig. 86). In Tipuloidea reduction of the hypostoma seems to be linked with reduction of the head capsule. Tipuloidea without a hypostoma possess a reduced head capsule, whereas first and second level out- group taxa without a hypostoma do possess a com- pact head capsule. Therefore, an anteriorly toothed hypostoma as found in Tipuloidea with a compact OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea head capsule must be considered the groundplan condition of Tipuloidea. It is not yet clear whether the groundplan hypostoma is divided, as in lower Hexatominae (figs. 58, 59), or is complete as in all other Tipuloidea with a compact head capsule, ex- cept Pediciini. A transformation series from a com- plete hypostoma could include the following steps: in the first instance the posterior cleft evolved as a continuation of the anterior extension of the occipital foramen (character C), subsequently div- iding the hypostoma, as in Pediciini, lower Hexa- tominae and lower Eriopterinae, with loss of the anterior teeth in higher Eriopterinae and higher Hexatominae. The above transformation series together with the monophyly of the Pediciinae presumes that the divided hypostoma of Pediciini is an independent development. In most Pediciini the two hypos- tomal plates bear three apical teeth. These plates can be considered homologous to the lateral part of the hypostoma of U/a consisting also of three teeth in all species of which larvae are known. It must be concluded that in Pediciini the central labial part of the hypostoma is not present and was replaced by the push-button connection to keep the two lateral parts together. ATYPHOPHTHALMUS THAUMASTOPTERA LIMONIA ANTOCHA ORIMARGA ELLIPTERA RHIPIDIA LIBNOTES MET ALIMNOBIA GERANOMYIA DICRANOMYIA DISCOBOLA Fig. 8. Phylogeny of the Limoniinae. Characters of the major groups 1. Margins of the externo- and interno-lateralia heavily sclerotized 2. Frontoclypeal apotome reduced (figs. 14, 19) 3. Genae with deep incisions (figs. 14, 21, 26, 33) Head capsules of the Eriopterinae and Hexato- minae are characterized by the above characters, to be considered derived in comparison with the more primitive head, as in the genera Ula to Helius of 223 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 figure 4. A posteriorly rectangular head capsule is present in Pseudolimnophila (fig. 58), Cryptolabis (Hynes 1963: fig. 3), Cladura (fig. 26) and Chionea (fig. 21), and might belong to the groundplan con- dition of the Eriopterinae-Hexatominae lineage. Further reduction of the head capsule in both sub- families involves reduction of the amount of scle- rotization of the area between the heavily sclerot- ized margins, resulting in the so-called dissected type of head, consisting mainly of elongate rods (e. g. figs. 13, 68, 69; see also characters 49, 67, and Crisp & Lloyd 1954: p. 280-281, Lindner 1959: p. 223-230). 4. Spiracles of the pupae not protuberant In the first and second level outgroups, and in a number of other Nematocera, the lateral abdomi- nal spiracles of the pupae are protuberant. This character is present in a number of Eriopterinae (Alexander 1920: Erioptera, Ormosia, Symplecta, Gonomyia, Scleroprocta; Byers 1974: Idiognopho- myta) and Hexatominae (Alexander 1920: Pseudo- limnophila, Wood 1952: Conosia), but not in the other Tipuloidea, except Dactylolabis. It is not clear whether the pupal spiracles are always protuberant in Hexatominae and Eriopterinae, but information about this character is not available for the majority of the taxa belonging to these two subfamilies. 5. Ventral anterior margins of genae slightly ob- lique or straight In the first and second level outgroups, but also in the majority of the other families of Nematocera and in Nannochoristidae the ventral anterior mar- gins of the genae which border the maxillae have a distinct oblique orientation, as in Eriopterinae (fig. 15) and Hexatominae (fig. 58). In all other Tipuloidea the orientation of these margins is slightly oblique (fig. 111) or straight (e. g. fig. 94). In Mecoptera (Nannochoristidae excluded), the ventral part of the head capsule is largely closed and difficult to compare but the ventral margins of the genae which border the maxilla are straight in Panorpa (Steiner 1930). 6. Prementum with protruding lateral corners In Pediciinae the prementum is not rounded anteriorly as in other Tipuloidea (character 16), but appears as a flattened plate with protruding lateral lobes and is apically set with hairs (Ula) or spat- ulate setae (Pediciini, fig. 103). The prementum of Pediciinae is not very similar to the prementum in the first, second and third level outgroups which frequently possess a twofold or bilobed prementum (Steiner 1930, Anthon 1943a). It is nevertheless possible to consider the somewhat bilobed pre- mentum of Pediciinae as symplesiomorphous with 224 respect to the anteriorly rounded prementum of the other Tipuloidea (character 16). 7. Larval antenna with two distinct papillae In the first and second level outgroups the larval antenna is usually very short. In Tipuloidea the antenna is usually elongated with a well developed basal segment, apically provided with small sen- silla, a few hairs, and one distinct papilla (e. g. figs. 19, 26, 76). Relatively short antennae are found in Ula (fig. 114) and some Dactylolabis species. In most Pediciini and Ula the basal segment of the antenna bears two distinct papillae (figs. 98, 99, 114). This synapomorphy must be considered weak. One papilla is present in Trichocera and most second and third level outgroups but two papillae are present in some Anisopodidae (Keilin & Tate 1940, Anthon 1943a). According to Alex- ander (1920), two papillae are also present in Pseu- dolimnophila but this apparently applies to P. in- ornata only (Alexander 1920: fig. 174) and not to P. luteipennis (Malloch 1917) and P. lucorum (fig. 76; Brindle 1960, Rozkosny & Pokorny 1980). Furthermore, the character is lacking in some Pe- diciinae (see character 97). 8. Posterior incisions of genae narrow or hardly developed Pediciini show the most massive head capsule. It is elongate without distinctly recognizable sutures and the posterior incisions of the genae are hardly developed (figs. 93, 94). A similar type of elongate head is present in Ula mollissima but with distinct frontal sutures (fig. 110). The absence of clearly developed posterior incisions of the genae in these taxa could be interpreted as a symplesiomorphy. However, a trend towards reduction of the poste- rior incisions is seen in other species of Ula, such as U. sylvatica and U. elegans, where the posterior incisions of the genae are well developed but nar- row (fig. 116). Distinct posterior incisions are present in the younger larvae of Ula and Pediciini as well (figs. 95, 96; see also Lindner 1959). 9. Pupae in a silken cocoon Larvae of Pediciini, U/a and Limoniinae con- struct a silken cocoon for the pupae. Among other Nematocera this character seems to be present only in Mycetophilidae (s.1.) and Simuliidae. On the basis of the adopted phylogeny it must be con- cluded that this character developed independently in Pediciinae and Limoniinae. Information about this character is not available for the limoniine genera Libnotes and Discobola (see also character 34). Larvae of the pediciine genus Dicranota living in lotic water apparently do not construct the pupal tubes unless they have some means of reaching the OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea | O, Figs. 9-10. Trichocera spec., left mandible: 9, ventral view; 10, lateral view. - Figs. 11-20. Eriopterinae, larval characters. - 11, Rhypholophus haemorrhoidalis, left mandible, lateral view; 12, idem, labial region, ventral view; 13, Molophilus spec., head capsule, ventral view; 14, R. haemorrhoidalis, head capsule (fc, frontoclypeal apotome; cs, coronal suture; ilp, elp, interno- and externo-lateral parts of genae), dorsal view; 15, idem, ventral view; 16, Gnophomyia jacobsoni, anterior ends of ventral bars with hyaline connection; 17, Cladura flavoferruginea, left mandible, lateral view; 18, Cheslotrichta cinerascens, hy popharyngeal bar; 19, R. haemorrhoidalis, anterior part of head capsule, dorsal view; 20, Chionea stoneana, larva (probably third instar), lateral view (figs. 9-10: Anthon 1943a; 11, 12, 14, 15, 19: Lindner 1959; 13: Alexander & Byers 1981; 16: Rogers 1927b; 17: Alexander 1922; 18: Reusch 1988; 20: Byers 1983) (all figures redrawn). 225 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 banks of the river or stream, so that occasional pupae may be found free (Brindle 1967). 10. Mandibles operating in a horizontal plane In the first and second level outgroups the mand- ibles have an oblique or vertical orientation, as in Eriopterinae, Hexatominae and Ula (Perris 1847, Alexander 1915a, own observation). The mandi- bles of Pediciini have a slightly oblique orientation while feeding (Wardle 1926), but apparently move in a much more vertical direction when used as an aid in moving forward (Wardle & Taylor 1926). Propulsion aided by the mandibles is also known for Ula (Perris 1847, Alexander 1915a, 1920). In the other Tipuloidea the mandibles operate horizontally, except in Cylindrotominae where they move vertical in second stage to full-grown larva, but horizontal in first stage larva (Bengtsson 1897, Alexander 1920, Peus 1952). Such a shift in the orientation of the mandibles is not known for other Tipuloidea and, for example, Byers (1983) recorded an oblique orientation in first instar to full grown larvae of Chionea (Eriopterinae). 11. Five-toothed hypostoma In Austrolimnophila, Epiphragma, Dicranopty- cha, Lipsothrix and first stage larvae of Tipulidae, the hypostoma is strikingly similar, consisting of three large central teeth in front and one small lateral tooth on either side situated more dorsally (figs. 124, 131). A five-toothed hypostoma is also present in Helius (fig. 135), Limnophilomyia and Elephantomyia aurantiaca (fig. 170). The presence of a five-toothed hypostoma in this wide range of taxa indicates that this is the basic number of hyp- ostomal teeth for part of the Tipuloidea with a complete hypostoma. A higher number of teeth is found in Eriopteri- nae (Molophilus 8, Erioptera 10), Hexatominae (Paradelphomyia 8-10, Pseudolimnophila 14-16), Pediciinae (Pediciini 6-8, Ula 9), Dactylolabis (9- 11), Atarba (7), Limoniinae (usually 9-11, in Dicra- nomyia, Libnotes and Metalimnobia up to 17), Cy- lindrotominae (about 15) and Tipulidae (full grown larvae 7-9)!. 12. Maxillary palpus The maxillary palpus in first and second level outgroups is flush with the surface of the maxilla, recognizable only by a ring of chitin surrounding the sensilla (chapter 3: character 45). This type of maxillary palpus is found in Eriopterinae (fig. 23), Hexatominae, Pediciini (the two last-mentioned taxa with the outer maxillary lobe elongate and the sensilla situated near or at the apex, figs. 68, 100), Ula (fig. 113) and Dactylolabis. In other Tipuloidea the sclerotized ring surrounding the sensilla is cy- 226 lindrical and usually well developed (figs. 121, 151, 163, 178), but with a low profile in Helius (fig. 133) and several species of Limoniinae (fig. 152). 13. Labral sclerites The genera Austrolimnophila and Epiphragma share a number of larval and pupal characters, such as shape of the mandible, antenna, maxilla, hypo- pharynx, armature of pupal abdominal segments (Alexander 1920, Bruch 1939, Wood 1952, Lindner 1959, Brindle 1960). A distinct synapomorphy for both genera is the peculiar shape of the labral sclerites (figs. 126, 129). 14. Pupal armature One or more rows of spines are usually present along the posterior margin of the pupal abdominal segments in Tipuloidea (e. g. fig. 47) and a great deal of other Nematocera (Brauns 1954b). Such rows are absent in lower Eriopterinae (character 53), Dicranoptycha, Helius, Lipsothrix, Limnophi- lomyia (fig. 153), Limoniinae (fig. 157-159; Dis- cobola caesarea excepted ?, see Mik 1884), and Cylindrotominae (fig. 156), but are present in Ti- pulidae. In Hexatominae the spines can be dis- tinctly tubular (character 75). 15. Hypopharynx dentated 16. Prementum dentated Below the ventral floor of the pharynx, most taxa of the Helius-Tipulidae lineage are provided with three rows of teeth, namely the hypopharynx, prementum and hypostoma (fig. 149).The hypo- pharynx of Tipuloidea and first and second level outgroups (Anthon 1943a), is usually preserved as a hemispherical cushion or lobe, densely set with setae or minor prominences and supported by a U- or H-shaped, sclerotized bar (figs. 122, 132, 175, 176). Deviations from this groundplan condition are: higher Hexatominae: sclerotized bar trans- formed into the very typical hypopharyngeal ske- leton (character 67). Pediciini: central part of bar not sclerotized (Reusch 1988: fig. 38B). Ula: hypo- pharynx incorporated in the pharyngeal filter ap- paratus (details in Lindner 1959). Helius, Limno- philomyia, Limoniinae and Cylindrotominae (character 15): hypopharynx not hemispherical but sclerotized and distinctly dentated (figs. 147, ' Five teeth is mentioned for Longurio minusculus Alex- ander (Wood 1952), and Tipula (Trichotipula) orope- zoides Johnson (Alexander 1920). In both instances the prementum has three teeth. In Tipulidae the prementum of the first stage larvae has three teeth and that of the full grown larvae five teeth. Three premental teeth in the above two species might indicate that the described larvae were not full grown. OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 21-25. Eriopterinae, larval characters. - 21, Chionea stoneana, head capsule, dorsal view; 22, idem, left mandible, ventrolateral view; 23, idem, left maxilla (mp, maxillary palp), ventral view; 24, Idiognophomyia enniki, head capsule, lateral view; 25, idem, terminal segment, ventral view (figs. 21-23: Byers 1983; 24, 25: Byers 1974) (all figures redrawn). 148, 167, 168). This synapomorphy must be con- sidered weak because it is absent in Lipsothrix (Rogers & Byers 1956, Hynes 1965) and Tipulidae (fig. 176). The prementum of Tipuloidea is usually hardly sclerotized and anteriorly rounded, the ventral sur- face provided with hairs, bristles or spines (fig. 122), and bearing the rudimentary labial papillae (figs. 122, 148). In lower Hexatominae (Pseudolimnophila) the outer margin is set with minor teeth. In Pediciinae the prementum has the outer lobes produced (fig. 103; character 6). A sclerotized and distinctly dentated prementum (character 16) is present in the taxa of the Helius- Tipulidae lineage (figs. 147, 148, 167, 168, 175, 176), Limnophilomyia excepted (Wood 1952). Li- mited information is available for Lipsothrix and Limnophilomyia but the prementum is very sim- ilar in Helius, Limoniinae and Tipulidae, with the orifice of the salivary duct between prementum and hvpopharynx. In Cylindrotominae the orifice is sit- uated more in front, just below the outer row of teeth (figs. 167, 168), indicating that this row of teeth might not belong to the prementum but to the hypopharynx. 17. Interno-lateralia fused with frontoclypeal apotome In Lipsothrix, Limnophilomyia, Limoniinae, Cy- lindrotominae and Tipulidae the internal lateral parts of the genae are fused with the frontoclypeal apotome (fig. 140). In all other Tipuloidea with a complete hypostoma the interno-lateralia and the frontoclypeal apotome are separated by the frontal sutures (fig. 134). 18. Inner maxillary lobe with a separate sclerotized plate A separate sclerotized plate on the inner maxil- lary lobe between the cardo and the sclerotization along the outer margin of the lobe is present in Limoniinae, Cylindrotominae and Tipulidae (figs. 151, 152, 163, 173, 178) and not in other Tipuloidea (figs. 100, 113, 121). It is not known whether this character occurs in Lipsothrix and Limnophilo- myia, and whether one of the sclerotized plates on the inner maxillary lobe in Helius (fig. 133) repres- ents the same plate. 19. Externo-lateralia strengthened In Limoniinae, Cylindrotominae and Tipulidae the externo-lateralia are strengthened by distinct sclerotizations bordering the genae in front of the area of the growth lines (figs. 139, 140, 160, 173). It is not known whether this character is present in Lipsothrix and Limnophilomyia. 20. Spiracular field with two dorsal lobes Two dorsal lobes in addition to the dorso-lateral and ventro-lateral lobes of the spiracular field are present in Tipulidae and Cylindrotominae only (figs. 164, 177). This synapomorphy is considered weak because it is not clear whether the dorsal lobes in Cylindrotominae do in fact represent ab- 224]; TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 dominal lobes. Furthermore, absence of these lobes in species of Cylindrotoma could represent the groundplan condition in Cylindrotominae. 21. Premaxillary suture and side plates In Tipulidae and Cylindrotominae distinct side plates are found (Peus 1952: ‘Seitenplatten’), se- parated from the ventral and lateral parts of the head capsule by a premaxillary suture sensu Cook (1949) (figs. 160, 173). In both families the side plates include the stemmata (Cook 1949, Peus 1952). In the other Tipuloidea the side plates and premaxillary suture are absent. 22. Prostheca on a separate sclerite A prostheca in the form of a brush of long hairs arising directly from the surface the mandible is found in all Nematocera (figs. 9-11). As far as is known only in full grown larvae of Tipulidae and Cylindrotominae the prosthecal brush is situated on a separate sclerotized lobe (figs. 162, 179, 180; see also chapter 3, Tipuloidea and ‘all other Nemat- ocera’ as sistergroups). 23. Hypostoma of first stage larvae different As outlined above, the presence of an anteriorly toothed hypostoma is considered to belong to the groundplan condition in Tipuloidea (character G). Several authors are of the opinion that the hypos- toma is of genal origin (e.g. Anthon 1943a, 1944b, Snodgrass 1960, Matsuda 1965), whereas others claim a labial origin (e.g. de Meijere 1917, Cook 1949, Saether 1971, see also Hennig 1973, Teskey 1981). The third alternative of a combined genal and labial origin, postulated by Bengtsson (1897) and again by Courtney (1990, 1991), is demon- strated in Cylindrotominae and Tipulidae (and might belong to the groundplan condition in Tipu- loidea with a complete hypostoma or even all Ti- puloidea), where the anterior part, the true hypos- toma, is of labial origin and the posterior part, the hypostomal or subgenal bridge, is of genal origin. In Tipulidae and Cylindrotominae there are marked differences between the hypostoma of the first stage larvae and the full grown larvae, whereas in other Tipuloidea the hypostoma of the first stage larvae and the full grown larvae seem in principal to be the same. The latter statement, however, is based on limited information. Availa- ble for examination were first stage larvae of Cy- lindrotominae (Diogma, Phalacrocera), Tipulidae, Limoniinae, (Limonia, Metalimnobia, Dicrano- myia), Helius, Austrolimnophila (Limnophilella), Hexatoma and Erioptera. Published accounts on the hypostoma of first stage larvae do hardly exist. To be mentioned are Bengtsson (1897, Cylindroto- minae: Phalacrocera replicata), Crisp & Lloyd 228 (1954, Tipulidae), and Okely (1979, Pediciinae: Tricyphona immaculata; Eriopterinae: Erioconopa trivialis). The hypostoma in first stage larvae of Tipulidae appears as a large, 5-toothed plate (see character 11). This plate lies in isolation behind the two sclerotized plates which represent the side plates. The isolation of the hypostoma from the genae indicates that it is of labial origin. In the full grown larvae of Tipulidae the general appearance of the hypostoma is the same as in the other Tipuloidea with a complete undivided hypostoma, and has become a 7- or 9-toothed plate, fused with the inner ventral margins of the genae (fig. 173). First stage larvae of Cylindrotominae (Diogma, Phalacrocera) (fig. 165) possess a complete sub- genal bridge which is maintained in the full grown larvae. Its presence can be considered plesiomor- phic (Teskey 1981), but in other Nematocera this bridge is formed by the posterior margin of the genae, whereas in Cylindrotominae it is formed anteriorly and homologous with the side plates. Situated in front of and slightly above this heavily sclerotized bridge, the first stage larvae possesses a small, completely hyaline, bilobed plate, the hy- postoma. Its position and the difference in sclerot- ization indicate that this small plate is of labial origin, as was pointed out already by Bengtsson (1897). In full grown larvae of Cylindrotominae the hypostoma has become a distinct, sclerotized and multidentated plate, completely fused with the subgenal bridge, but with distinct lines of fusion (fig. 166). As far as is known, the hypostoma of the first stage larvae of the other Tipuloidea does not show the developments found in Tipulidae and Cylindro- tominae. In the first stage larvae of the other Ti- puloidea the hypostoma already possesses the same number of teeth as found in the full grown larvae, and it is fused with the ventral margins of the genae. In several genera a distinct line is visible at the transition of the genae and the hypostoma, apparently representing the line of fusion. It must be concluded that in the other Tipuloidea fusion of genae and hypostoma already occurs in the egg. 24. Head capsule elongate In comparison to Mecoptera, Nematocera and other Tipuloidea (e. g. fig. 119), the Eriopterinae, Hexatominae and Pediciinae possess an elongated head capsule, about twice as long as broad (e. g. figs. 14, 66, 93). These elongate head capsules are char- acterized by the very narrow coronal suture. 25. Interno-lateralia extended dorsally In larvae of Eriopterinae, Hexatominae and Pe- diciini the interno-lateralia of the genae occupy OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea 37 È yo —- Figs. 26-37. Eriopterinae, larval characters. - 26, Cladura flavoferruginea, head capsule (lp, labral papilla), ventral view; 27, Baeoura claripennis, terminal segment, dorsal view; 28, Cryptolabis magnistyla, terminal segment, dorsal view; 29, idem, lateral view; 30, Cheilotrichia cinerascens, spiracular disc; 31, Molophilus spec., spiracular disc; 32, Chionea stoneana, terminal segment, posterodorsal view; 33, Gonomyia sulphurelloides, head capsule, dorsal view; 34, Eugnophomyia luctuosa, spiracular disc; 35, Ellipteroides lateralis, spiracular disc; 36, Scleroprocta spec., larva, dorsal view; 37, Idiognophomyia enniki, larva, lateral view (fig. 26: Alexander 1922; 27, 33: Wood 1952; 28, 29: Hynes 1963; 30, 31: Lindner 1959; 32: Byers 1983; 34: Rogers 1928; 35: Bangerter 1928; 36: Alexander 1920; 37: Byers 1974) (all figures redrawn). 229 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 most of the dorsal part of the head capsule and the frontoclypeal apotome is reduced (fig. 14) or en- tirely fused with the interno-lateralia (fig. 93). 26. Penultimate segment inflated Larvae of Eriopterinae, Hexatominae and Pedi- ciini are able to inflate the penultimate segment (fig. 64). This character, not known from first to third level outgroups or other Tipuloidea except Atarba picticornis (Rogers 1927a), is widespread in all three taxa but is not recorded for all genera, most probably because it is not a permanent, struc- tural character. Inflation is observed most fre- quently in moving animals and is achieved within a few seconds, apparently to afford anchorage to the substrate (Crisp & Lloyd 1954, Lindner 1959). 27. Outer maxillary lobe lengthened The maxillary lobes of Tipuloidea larvae are usually short (e. g. figs. 23, 121) as in other Nema- tocera, whereas in Hexatominae, Pediciini and some Eriopterinae (Cryptolabis and Rhabdomas- tix) the outer maxillary lobe is lengthened (figs. 58, 60, 66-69, 100). Lengthened maxillae are usually considered to be correlated with the predatory feeding of Hexatominae and Pediciini but they are also present in the less- or non-predatory genera Paradelphomyia, Pseudolimnophila, Neolimno- myta, Cryptolabis and Rhabdomastix. The pres- ence of lengthened maxilla in especially the two last-mentioned and apparently not closely related genera indicates, that this character evolved inde- pendently in Eriopterinae and perhaps also in Hexatominae and Pediciini. The outer maxillary lobe of Pseudolimnophila (fig. 58) with two joints, each with a ring sclerite, very much resembles that of Mecoptera (Steiner 1930), including Nannochoristidae (Pilgrim 1972), but in Pseudolimnophila the inner lobe is not dis- tinctly separated from the outer lobe (Crisp & Lloyd 1954). 28. Spiracles on dorsum of pupal tergite 8 Nematocera pupae apparently do not possess spiracles on the dorsum of tergite 8. An account of such spiracles was found only for the Pachyneuri- dae (Krivosheina & Mamaev 1970). In Tipuloidea spiracles on the dorsum of tergite 8 are in general absent in Eriopterinae, Hexatominae, Dactylolabis and Dicranoptycha, but are present in most of the other groups except Cylindrotominae and Tipuli- dae, of which the pupae are propneustic although some lower Tipulidae do possess spiracles on the dorsum of tergite 8 (Wood 1952). It is not known whether this character occurs in Limnophilomyia, Lipsothrix, and several Limoniinae (Discobola, Ge- ranomyia, Rhipidia). In Hexatominae these spira- 230 cles are present in Paradelphomyia senilis (Reusch 1988), Euphylidorea meigeni (own observation) and they are figured by Wiedenska (1987) for Phy- lidorea nigronotata but are absent in the other Phylidorea species studied by her. In Limoniinae they are absent in Antocha and Elliptera (own observation). 29. Larvae without ventral creeping welts Presence of ventral creeping welts, densely set with recurved hooklets or spinules, can be consi- dered plesiomorphic. Such welts are found in lar- vae of many Nematocera and Tipuloidea. In Tipu- loidea they are in general present on the anterior part of segments 5-10 (e. g. figs. 37, 109). Some Limoniinae possess additional creeping welts on segments 2-4 and 10 (figs. 143, 144). In Trichocera ventral creeping welts are not present. In Eriopte- rinae and Hexatominae presence of ventral creep- ing welts, again on sternites 5-10, is restricted to a few genera (characters 48, 79). In the Pediciini ventral creeping welts are restricted to sternites 6- 10 or 7-10 (figs. 107, 108), with the welts deve- loped into pseudopods in certain genera (fig. 104; characters 84, 91, 95). Several species of Epiph- ragma and certain subgenera of Tipula (Gelhaus 1986: Arctotipula, Nippotipula, Sinotipula) have naked transverse swellings which lack recurved hooklets or spinules. Ventral creeping welts are absent in Dactylolabis and Cylindrotominae. 30. Larvae with dorsal creeping welts Presence of dorsal creeping welts can be consi- dered derived because Nematocera larvae rarely possess dorsal creeping welts (in contrast to Bra- chycera). Dorsal creeping welts occur frequently again on the anterior part of segments 5-10, with additional welts in certain Limoniinae. In Trichoc- era dorsal welts are absent. Among the higher Hexatominae dorsal creeping welts on tergites 5- 10 are restricted to a few genera (character 79). Dorsal creeping welts in Pediciini are known for Rhaphidolabina flaveola only (Alexander 1920). A very special arrangement of convergent dorsal welts is found in a number of species of Dactylo- labis (Nowicki 1867, Bangerter 1931, Sinclair 1988). In Tipulidae naked transverse dorsal swel- lings are present in Tipula (Nippotipula) (Gelhaus 1986). 31. Pupae with ventral creeping welts 32. Pupae with dorsal creeping welts Pupae of Tipuloidea frequently possess dorsal and/or ventral creeping welts on the abdominal segments (figs. 153, 154, 157). It is not certain whether presence of these welts should be consi- dered derived, but welts seem to be rare in pupae OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea | i WX RS Ÿ SE sk As, MS ee Mr ARRET Figs. 38-44. Eriopterinae, larval characters. — 38, Scleroprocta spec, terminal segment, ventral view; 39, Rhypholophus haemorrhoidalis, spiracular disc; 40, Gnophomyia toschiae, terminal segment, oblique posterior view; 41, Gonomyia spec., terminal segment, oblique posterior view; 42, Erioconopa trivialis, spiracular disc of first instar; 43, Rhabdomastix californiensis, terminal segment, oblique posterior view; 44, Ormosia spec., terminal segment, oblique posterior view (fig. 38: Alexander 1920; 39: Lindner 1959; 40, 41, 44: Alexander & Byers 1981; 42: Okely 1979; 43: Hynes 1969c) (all figures redrawn). 291 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 of other Nematocera (Brauns 1954b). In pupae of Trichocera welts are absent. In pupae of Tipuloidea the welts usually occur on abdominal tergites 3-7 and sternites 5-7. In the Eriopterinae dorsal and ventral welts are present in the pupae of Gropho- myta, Idiognophomyia, Hesperoconopa, and some species of Gonomyia. The creeping welts in pupae of Pediciini are discussed under character 96. 33. Pupae with inner legsheaths the longest In Mecoptera including Nannochoristidae and the majority of the Nematocera and Tipuloidea, the outer legsheaths of the pupae are the longest (fig. 154) although in Eriopterinae, Hexatominae, Epiphragma, Ula and Cylindrotominae, some spe- cies or specimens do possess leg sheaths of equal length. In Dactylolabis, Helius, Limnophilomyia and Limoniinae the inner legsheaths are usually the longest but also in these taxa, species or spec- imens might show equal leg sheaths. Mik (1886) figured outer leg sheaths the longest in Elliptera, but pupae of this genus examined had equal leg sheaths. The special arrangement of the leg sheaths in Eriopterinae is discussed under charac- ter 34. Larvae with spinning glands The larvae of a number of species possess spin- ning glands. Except for Cylindrotoma, the larva of the species concerned constructs a case or a cocoon in which pupation takes place. In Dicranoptycha and the limoniine genus Thaumastoptera a case is constructed out of small pieces of wood, leaves or fine grains of sand. All species with spinning glands, except Dicranoptycha, possess real silk glands to produce a silken cocoon covered with detritus for pupation or, in Thaumastoptera, to produce a silken inner lining to their cases. Many larvae of Limoniinae are living in silken tubes themselves as well (character 99). The only account for Cylindrotominae is by Cameron (1918) who reported for the terrestrial Cylindrotoma splendens |= distinctissima americana | that young larvae have the power ‘of secreting a silken thread from the mouth, which is probably the product of the salivary glands. They frequently adhered to this thread ... used in transferring them from one leaf to another’. 35. Apophyses present A number of Tipuloidea possess small sclerot- ized appendages posterior to the hypostoma (figs. 97, 135). They play part in the attachment of the prothoracic skin to the larval head capsule (Peus 1952), and are not present in other Nematocera or the related orders. Apophyses are not figured or described for Dactylolabis (Bangerter 1931, 232 Brindle 1960, Sinclair 1988), but in Dactylolabis montana they are present (own observation). 5. ERIOPTERINAE Introduction The monophyly of the Eriopterinae is based on two rather weak characters only (fig. 5). Larvae of Eriopterinae usually possess a narrow labrum (figs. 13, 19, 21, 26, 33), slender mandibles (figs. 11, 17, 22), simple maxillae (fig. 23) and relatively well developed antennae with a distinct, usually some- what asymmetrical apical papilla (figs. 19, 21, 26). These characters are not listed as synapomorphies because they might as well represent the ground- plan condition in Tipuloidea. The amount of information on the head capsule of Baeoura is limited, described by Wood (1952) for the species claripennis as follows: ‘Head capsule long and slender, consisting of three concave plates, one dorsal and two lateral, the margins of which are heavily chitinized. Labrum and maxilla as in the genus Gonomyia. Remainder of head capsule difficult to distinguish and cannot be accu- rately described’. Character 49, head capsule further reduced, is taken as landmark between the lower and higher Eriopterinae. The lower Eriopterinae include the Cladurini (the genera Cryptolabis, Cladura and Chionea). Character 39 suggests that this tribe can also be recognized on the basis of the larvae. Genera belonging to the Gonomyiini are found in one section of fig. 5 except Gonymyodes, partly as lower and partly as higher Eriopterinae. Styringo- myia, placed in the Eriopterini by Hynes (1990) is found amidst the Gonomyiini. The tribes Eriopte- rini and Molophilini do not show a close corres- pondence to the phylogenetic arrangement pres- ented. Eriopterinae larvae are gross-feeding forms on decayed plant remains (Crisp & Lloyd 1954). Strictly predatory forms are not known, but accord- ing to Byers (1983), Chionea certainly is not a typical herbivorous tipuloid and might be carnivor- ous. In the text below reference is made to characters of the genera included in fig. 5. Other eriopterine genera are mentioned in chapter 9. Characters 36. Last larval segment constricted In almost all Eriopterinae the last larval segment is basally constricted (figs. 36, 37) and can be tel- escoped into the penultimate segment during pro- gression, a way of propulsion also recorded by Alexander (1920) for Ula. No constriction is found in Cladura and Chionea (fig. 20). Information about this character is equivocal or not available for Eugnophomyia, Arctoconopa, Erioconopa, Ilisia and Hoplolabis. This synapomorphy must be con- sidered weak because it is also found in a number of Hexatominae (character 76). 37. Sheaths of middle legs the shortest Pupae of Eriopterinae species frequently have the middle legsheaths the shortest (figs. 45,52, 54), usually with the outer legsheaths the longest, but sometimes with the inner and outer legsheaths of equal length. This character is not present in other Tipuloidea except Atarba (see chapter 9) and ap- parently does not occur in other Nematocera. Therefore, it is listed here as a synapomorphy al- though it is not exhibited by all Eriopterinae. The character is present in Cryptolabis, Cladura, Styrin- gomyia ingrami (Edwards 1924), Eugnophomyia, Gonomyia, Ellipteroides, Cheilotrichia, Symplecta, Ormosia, Rbypholophus, Gonomyodes, Ilisia, Ho- plolabis, Erioptera and Molophilus. The legsheaths are of equal length in Baeoura (Wood 1952), some species of Grophomyia (Malloch 1917), in Styrin- gomyia neocaledoniae (Hynes 1990) and in Sclero- procta (Alexander 1920, Bangerter 1930, Brindle 1967). Outer legsheaths the longest and inner leg- sheaths the shortest is found in Chionea (Byers 1983), Idiognophomyia (Byers 1974), Gnopho- myia (Alexander 1920; Rogers 1927b) and Rhab- domastix (Hynes 1969c). Information about this character is not available for Arctoconopa and Eri- oconopa. 38. Pupae without respiratory horns Pupae of Tipuloidea usually possess respiratory horns, present also in all Eriopterinae included in fig. 5, except Baeoura, Cryptolabis, Cladura, and Chionea. 39. Labral papilla Larvae of Cryptolabis, Cladura (fig. 26), and pos- sibly also Chionea (fig. 21) possess ‘two large cy- lindrical papillae which arise near the clypeo-labral suture and which appear to belong to the labrum’ (Hynes 1963 for Cryptolabis), not present in other Tipuloidea or Nematocera. 40. Last larval segment not constricted Synapomorphy 36 is not present in Cladura and Chionea (fig. 20). 41. Spiracular lobes absent Cladura and Chionea (fig. 32) do not possess spiracular lobes (see also character 43). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea 42. Palpsheaths obliquely directed downward In Psychodidae, Trichoceridae, Anisopodidae, Scatopsidae and most other families of Nematoc- era, the palpsheaths of the pupae are directed ob- liquely upward. This is also the case in the majority of the Tipuloidea with a compact head capsule. In Hexatominae the maxillary palps are usually horiz- ontal or apically curved downward (figs. 87-90). In Phylidorea, Euphylidorea, Eloeophila and Hexa- toma horizontal as well as slightly upcurved sheaths are found. In Eriopterinae the sheaths of the palps are obliquely directed downward (e. g. fig. 47), except for Baeoura (upward), Cryptolabis (ho- rizontal), Chionea (upward, fig. 46) and Cladura (upward). 43. Spiracular field with five more or less identical lobes 44. Darkened spots on ventral spiracular lobes divided into two The apparent plesiomorphic condition of four spiracular lobes is present in Baeoura (fig. 27) and Cryptolabis (figs. 28, 29). In these two genera the lobes are rounded and without dark markings. Lobes are absent in Cladura and Chionea (character 41). The other Eriopterinae usually possess five more or less identical lobes with the dark markings on the ventral lobes divided into two (e. g. fig. 39). The number of spiracular lobes is seven in Styrin- gomyia and Gonomyodes, and nine in Ellipteroides (fig. 35), while the genera Rhabdomastix, Arcto- conopa and Gonomyia have shortened lobes. Character 44 must be considered a weak synapo- morphy, because it is also present in a number of Hexatominae (character 80). It is not found in Styringomyia, Gonomyodes and some species of Rhabdomastix and Symplecta (see also characters 290162): 45. Pupae with dorsal and ventral creeping welts 46. Respiratory horns earshaped 47. Anterior anal papillae longer than posterior ones Idiognophomyia and Gnophomyia are consi- dered sister genera on the basis of the above three characters although character 45 is found in Gono- myia as well, character 46 in some species of Go- nomyia (fig. 48) and Ellipteroides, and character 47 (fig. 25) in Scleroprocta, Ormosia (fig. 44) and Rhypholophus. 48. Larvae with ventral creeping welts As outlined above (character 29), larval creeping welts on sternites 5-10 is apparently plesiomor- phic. This character is present in Cladura, Chionea (fig. 20), Idiognophomyia (fig. 37) and Gnopho- myta. Loss of creeping welts can be considered a 233 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 synapomorphy for the higher Eriopterinae but oc- curred also in Baeoura and Cryptolabis. Ventral creeping welts are absent as well in the majority of Hexatominae (character 79). Therefore, it appears more parsimonious to accept presence of ventral creeping welts as independent returns to the ple- siomorphic condition in the Chionea-Cladura and Idiognophomyia- Gnophomyia lineages. 49. Head capsule further reduced A still rather compact head capsule is found in the so-called lower Eriopterinae (figs. 21, 26). In the higher Eriopterinae the head capsule is reduced further. The amount of sclerotization between the heavily chitinized margins is minimized and the head capsule appears as consisting mainly of elon- gate, strongly sclerotized rods (figs. 13-15, 33). These rods are homologous to the heavily chitin- ized margins which border the frontoclypeal apo- tome and loop around the genae. A break in the loop around the genae is found already in Idzog- nophmyia and Gnophomyia (fig. 24). 50. Mesothorax of pupae with distinct crest Pupae of higher Eriopterinae, Styringomyia and Gonomyodes excepted, possess a distinct crest on the mesothorax (figs. 47, 48, 50, 51). This crest is absent in other Tipuloidea, some South African species of Limnophila excepted (fig. 87; Wood 1952). 51. Larval antennae near to each other In higher Eriopterinae, Styringomyia excepted, the larval antennae are situated very near to each other (fig. 19). This character is not found in other Tipuloidea except Idiognophomyia. No informa- tion is available for Baeoura, Rhabdomastix and Gonomyodes. 52. Central part of spiracular field with a distinct dark marking Among Eriopterinae a distinct dark marking be- tween the spiracles is present in Ewgnophomyia, Gonomyia and Ellipteroides only (figs. 34, 35, 41). 53. Pupal abdominal spines Distinct rows of spines (figs. 47, 49, 50) are usually present along the posterior margin of the abdominal segments of the higher Eriopterinae, except for Styringomyia, Rhabdomastix Eugno- phomyia, Gonomyia and Ellipteroides, whereas no information about this character is available for Arctoconopa and Erioconopa. Similar rows of spines are present in other Tipuloidea (character 14). Their absence in lower Eriopterinae might indicate that spines in higher Eriopterinae is a de novo development. 234 54. Larvae with statocysts Statocysts, lateral sacs on the last abdominal seg- ments of the larvae, have been observed in species of Cheilotrichia, Symplecta, Ormosia, Rhypholo- phus, lista, Erioptera and Molophilus (Wolff 1922; Crisp & Lloyd 1954). Information for most other Eriopterinae is not available, but statocysts are ap- parently absent in Chionea (Byers 1983), Gono- myta (Crisp & Lloyd 1954) and all other Nematoc- era and Tipuloidea except for some higher Hexatominae (Wolff 1922, Crisp & Lloyd 1954; character 68). A detailed account of these organs is given by Wolff (1922), their functioning as statocysts is des- cribed by von Studnitz (1932). Statocysts control the geotactic reactions and are thought to be asso- ciated with larvae that swiftly move through the mud (Crisp & Lloyd 1954). 55. Hairs on spiracular lobes In Eriopterinae with five or more spiracular lobes, the hairfringe around the spiracular field is usually continuous (figs. 30, 32, 34, 35, 40, 41). In the Symplecta-Molophilus lineage the hairfringe is interrupted and the hairs are concentrated on the spiracular lobes (figs. 31, 39, 44). This character must be considered weak because the original con- figuration of the hairfringe is not always main- tained in preserved material, and likewise not al- ways accurately described or illustrated. Furthermore, this character is apparently present in Idiognophomyia (Byers 1974) and some species of Rhabdomastix. Crisp & Lloyd (1954) make the observation that, when the division of the dark markings (characters 44, 61, 62) is pronounced, the fringes are restricted to the tips of the spiracular lobes, whereas a slight division of the markings is associated with a com- plete hairfringe, representing the primitive condi- tion. 56. Shape of mandible Ormosia and Rhypholophus are characterized by the identical shape of the mandibles (fig. 11). Sim- ilar shapes are not found among Tipuloidea but are present, as bifold mandibles, in several Psychodo- morpha sensu Wood & Borkent 1989 (fig. 10). Ormosia and Rhypholophus do not represent the basal lineages of Eriopterinae. Therefore, the oc- currence of similar mandibles in these two genera and in Psychodomorpha must be considered a con- vergent development. 57. Dark marking along inner margin of spiracles In Arctoconopa, Erioconopa, Ilisia, Hoplolabis, Erioptera and Molophilus the spiracular field is provided with a small dark marking along the OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea 48 ns ERREUR EEE 3 PAS Ms sar) % 5 NEGEN Ten bye AN omega } 50 Figs. 45-56. Eriopterinae, pupal characters. - 45, Eugnophomyia luctuosa, female, ventral view; 46, Chionea lutescens, female, lateral view; 47, Erioptera chlorophylla, female, lateral view; 48, Gonomyia alexanderi, male, lateral view; 49, Ormosia nigripila, male cauda, dorsal view; 50, idem, male, lateral view; 51, Gonomyia sulphurella, female, lateral view; 52, Teucholabis complexa, male, ventral view; 53, longitudinal section through pupa of Erioptera squalida, showing insertion of respiratory horn into the gas space of a Glyceria root; 54, Erioptera cladophoroides, tarsal sheaths; 55, Symplecta pilipes, male cauda, dorsal view; 56, idem, female cauda, dorsal view (fig. 45: Rogers 1928; 46: Byers 1983; 47-52: Alexander 1920; 53: Houlihan 1969; 54: Bruch 1939; 55, 56: Wood 1952) (all figures redrawn). 235 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 57-65. Hexatominae, larval characters. - 57, Pseudolimnophila inornata, terminal segment, oblique posterior view; 58, idem, head capsule (ml, outer maxillary lobe), ventral view; 59, Paradelphomyia minuta (supposition), hypostoma, ventral view; 60, idem, maxilla; 61, Ulomorpha spec., right mandible (pr, prostheca), dorsal view; 62, Eloeophila maculata, labial palp; 63, idem, hypopharyngeal skeleton, ventral view; 64, Hexatoma spec., larva, dorsal view; 65, Limnophila larva employing the head to stroke the long hairs of the spiracular disc free from the tension of the water surface (figs. 57, 58: Alexander & Byers 1981; 59, 60: Alexander 1920; 61: Anthon 1943a; 62, 63: Pokorny 1978; 64, 65: Lindner 1959) (all figures redrawn). 236 OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 66-69. Hexatominae, larval head capsules. - 66, Eutonia spec. (supposition), dorsal view, 67, idem, ventral view; 68, Conosia irrorata, ventral view, left dorsal bar interrupted to show hypopharyngeal skeleton, tip of right maxilla enlarged to show maxillary palp; 69, Limnophila crepuscula, dorsal view (Figs. 66, 67: Hennig 1950; 68, 69: Wood 1952) (all figures redrawn). 237 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 inner margin of the spiracles (fig. 31), not present in other Eriopterinae. 58. Ventral spiracular lobe without seta The ventral spiracular lobes are usually provided with a distinct seta in most Tipuloidea (e.g. figs. 44, 136), but not in Gonomyodes, Erioconopa, Ilisia, Hoplolabis, Erioptera and most species of Molo- philus. 59. Ventral rods of head capsule not apically dentated Hexatominae and Eriopterinae do not possess a complete hypostoma, but the ventral margins of the genae are still apically dentated in the lower representatives of these subfamilies (fig. 26). In the higher Eriopterinae the ventral rods of the head capsule (which are homologous with the ventral margins of the genae) are usually not dentated (fig. 12) except for some species of Rhabdomastix (Hynes 1969c: fig. 8) and for Erioptera and Molo- philus (fig. 13), which possess a similar hypos- toma, not found in other Eriopterinae. Therefore, return to the plesiomorphic condition is inter- preted as a synapomorphy for the two last-menti- oned genera. 60. Penultimate segment of larvae inflated Larvae of several Eriopterinae are recorded to be able to inflate the penultimate segment (fig. 36). Inflation is usually less conspicuous, although Wood (1952) records distinct inflation as in Hex- atominae for Rhabdomastix afra (see also charac- ter 26). 61. Dorsal spiracular marking divided into two 62. Lateral spiracular markings divided into two Apart from divided ventral markings (character 44), the markings on the dorsal and/or lateral spiracular lobes can be divided into two as well. 6. HEXATOMINAE Introduction The name Hexatominae is used here in a more restricted sense than in the current systematic liter- ature (chapter 4, Introduction). The genera Para- delphomyia and Pseudolimnophila are treated here as lower Hexatominae and all other genera of which larvae and/or pupae are known as higher Hexatominae. Paradelphomyia is usually assigned to the small tribe Paradelphomyiini, whereas Pseu- dolimnophila is included in the large tribe Limno- philini. In Hexatominae the spiracular field usually pos- sesses four elongate flattened lobes with the ven- 238 tral ones the longest, sometimes with a vestigial fifth lobe. The spiracular field is usually provided with long hairs, especially towards the tips of the elongate ventral lobes (figs. 57, 70, 71). By means of these long hair fringes the larva can hold air when submerged, or be freely suspended. The ef- fectiveness of holding the spiracular disc at the surface of the water can be so great, that the larva may have to curl over the spiracular disc in order to break the suspension (fig. 65). Four elongate, cylindrical instead of flattened lobes with the ventral ones the largest are also present in Cryptolabis (figs. 28, 29) and more or less in Baeoura (fig. 27). According to Hynes (1963), presence of elongate spiracular lobes in Cryptolabis and, for example, Eloeophila must be considered an adaptation to the habitat of the lar- vae, which live in sand and gravel of swiftly flowing waters. Elongate flattened spiracular lobes with the ventral lobes the longest is nevertheless not listed as a synapomorphy for the Hexatominae because it might as well be considered a synapomorphy for Eriopterinae plus Hexatominae. No other charac- ters are available to base upon the monophyly of the Hexatominae. The very similar hypostomal plates of Paradel- phomyia (fig. 59) and Pseudolimnophila (fig. 58) are not found among other Tipuloidea, but the similarity might be symplesiomorphic (see charac- ter G). Most Hexatominae are predators on insect lar- vae, oligochaetes, copepods and the like. Their me- thod of feeding is discussed by Crisp & Lloyd (1954) and a detailed description of a 10 mm Limnophila larva engulfing a 15 mm oligochaete is given by Lindner (1959). Pseudolimnophila is a gross feeder of plant remains, diatoms and small algae. Species of Paradelphomyia and Neolimnomyia are be- lieved to be feeders on animal protein, but are not always predacious or carnivorous (Crisp & Lloyd 1954, Hinton 1954, Brindle 1967). Characters 63. Setae in front of anal papillae In Hexatominae the last abdominal segment of the larvae ventrally bears distinct setae in front of the anal papillae, either arranged in groups (Wolff 1922), or as a distinct row (figs. 70, 71, 79-81). Information about these setae, which are absent in other Tipuloidea except for some species of Ele- phantomyia (see chapter 9), is not available for Paradelphomyia, Polymera and Neolimnomyia. 64. Tufts of hair on labrum The labrum of larvae of Hexatominae, Paradel- phomyia excepted, is provided with distinct tufts of OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 70-77. Hexatominae, larval characters. - 70, Ewtonia spec. (supposition), terminal segment, lateral view; 71, Conosia irrorata, terminal segment, lateral view; 72, Neolimnomyia nemoralis, spiracular disc; 73, Eloeophila sub- marmorata, labrum, dorsal view, showing large sclerotized plates on ventral surface caudal to epipharyngeal bar (eb); 74, E. dubiosa, labrum, dorsal view; 75, Conosia irrorata, labrum, dorsal view; 76, Pseudolimnophila lucorum, antenna; 77, Hexatoma cinerea, left hypopharyngeal plate, ventral view (fig. 70: Hennig 1950; 71: Wood 1952; 72: Brindle & Bryce 1960; 73: Pokorny 1978; 74, 75: Wood 1952; 76: Rozkosny & Pokorny 1980; 77: Alexander 1920) (all figures redrawn). hair on the antero-lateral sides (figs. 73-75). These tufts are not found in other Tipuloidea. It is not known whether they are present in Neolimnomyia or Eutonia. 65. Reduction of the inner maxillary lobe In Paradelphomyia two distinctly separated maxillary lobes are present (fig. 60). In the other Hexatominae there is a tendency to reduction of the inner lobe through fusion with the outer lobe (details in Crisp & Lloyd 1954, Pokorny 1978; re- duction of the inner lobe occurred also in several Pediciini, Cryptolabis and Rhabdomastix). 66. Mandible sickle shaped The mandibles of higher Hexatominae are sickle shaped with a long terminal point and shorter teeth about half-way down the blade (figs 61, 66- 69). A similar mandible is found in a number of Pediciini, probably as an adaptation to predation. The more generalized type of mandible, as in most other Tipuloidea including Paradelphomyia and Pseudolimnophila, usually has a strong row of teeth along the ventral edge and a weaker row along the dorsal edge (figs. 115, 125, 128, 138). 67. Head capsule further reduced The reduced head capsule of higher Hexatomi- nae does not show a dentated hypostoma divided along the midline as in Paradelphomyia and Pseu- dolimnophila, but the ventral rods possess medial side branches which articulate with the very typical hypopharyngeal skeleton. This skeleton consists of two large lateral plates and a transverse bar. This bar is perforated by an orifice and provided with very small teeth (figs. 63, 67-69). Furthermore, 239 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Va ALAS ent NW Wildy vili PALMA a a Figs. 78-85. Hexatominae, larval and pupal characters. - 78, Eutonia barbipes, spiracular disc; 79, Pseudolimnophila lucorum, terminal segment, ventral view; 80, Euphylidorea lineola, terminal segment, ventral view; 81, Pilaria discicollis, terminal segment, ventral view; 82, Limnophila fuscovaria, female cauda, lateral view; 83, idem, male cauda; 84, Neolimnomyia batava, terminal segments, lateral view; 85, Limnophila spec., terminal segments, oblique posterior view (fig. 78: Wolff 1922; 79-81: Savchenko 1986; 82, 83: Alexander 1920; 84: Brindle & Bryce 1960; 85: Alexander & Byers 1981) (all figures redrawn). well developed labial papillae are present (figs. 62, 63, 67). The hypopharyngeal skeleton is absent in Pilaria, Ulomorpha and several species of Hexa- toma whereas other species of Hexatoma possess large lateral plates (fig. 77) but lack the transverse bar. 68. Statocysts As in higher Eriopterinae, statocysts are present in larvae of higher Hexatominae. According to Wolff (1922) and Crisp & Lloyd (1954) they are not present in Paradelphomyia and Pseudolimnophila, while no information is available for Ulomorpha, Polymera, Neolimnomyia and Conosia (see also character 54). 69. Caudal end of dorsal plates spatulate 70. Mandible bifold Pilaria and Ulomorpha are without doubt sister- 240 groups on the basis of the above two characters. The very characteristic bifold mandibles (fig. 61) and the absence of any rigidity in the ventral region of the mouth are associated with the method of feeding on worms as described by Crisp & Lloyd (1954). 71. Maxilla narrowed towards tip The apex of the outer lobe of the maxillae is blunt in Paradelphomyia (fig. 60), Pseudolimno- phila (fig. 58), Pilaria and perhaps also Ulomor- pha. In the other Hexatominae the maxillae are narrowed towards the tips with a very characteris- tic outward orientation of the apical part (figs. 66- 69). 72. Dorsal bridge divided The dorsal plates of the head capsule are still fused in Pilaria and Ulomorpha (Alexander 1920) OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 86-92. Hexatominae, pupal characters. — 86, Conosia irrorata, male, lateral view; 87, Limnophila crepuscula, male, lateral view; 88, Ulomorpha pilosella, female, lateral view; 89, Eloeophila maculata, female, ventral view; 90, E. dubiosa, male, lateral view; 91, Pseudolimnophila lucorum, apical flaps of respiratory horn; 92, Conosia irrorata, respiratory horn (figs. 86, 87, 90, 92: Wood 1952; 88: Alexander 1920; 89: Pokorny 1978; 91: Hinton 1954) (all figures redrawn). but are separated and variously modified in the other higher Hexatominae (figs. 66, 68, 69). The description and figures by Rogers (1933) for Poly- mera can not be interpreted adequately but the dorsal plate is apparently divided (with large lat- eral plates as in Hexatoma ?, see Rogers 1933: fig. 2); In the Polymera-Eutonia lineage the anal papil- lae are short (figs. 64, 70, 71, 80, 84, 85) compared to the very elongate papillae of Paradelphomyia, Pseudolimnophila, Pilaria and Ulomorpha (figs. 57,79). This character might be considered a weak synapomorphy as well. Some species of Pilaria, however, can have the papillae less elongate (fig. 81), whereas they are elongate in Hexatoma spt- nosa (Byers 1978: fig. 19.17). 73. Apex of respiratory horn not flap-like In Pseudolimnophila, Pilaria, Ulomorpha and Polymera the respiratory horns are provided with apical flaps (figs. 88, 91). This character is inter- preted as belonging to the groundplan condition in Hexatominae except Paradelphomyia, with secon- dary loss in part of the higher Hexatominae (see also chapter 4, Respiratory horns). 74. Respiratory horn shortened In the Phylidorea-Eutonia lineage (figs. 86, 87, 89,90) the respiratory horns are much shorter than in Paradelphomyia and the genera with flap-like respiratory horns. The only exception are species of Neolimnomyia (Neolimnomyia) which also possess elongate respiratory horns. In Phylidorea, Euphylidorea, Eloeophila and several species of Limnophila the organ is laterally compressed. In some Hexatoma species they are acute (see also chapter 4: Respiratory horns.). 241 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 75. Pupal abdominal segment 7 with spined tuber- cles In a number of higher Hexatominae the seventh abdominal segment of the pupae is provided with many spined tubercles surrounding the caudal end of the segment (figs. 82, 83). It is not known whether this character is present in Polymera. It is apparently absent in Neolimnomyia. 76. Last abdominal segment constricted In Limnophila, Neolimnomyia, Conosia and Eu- tonia the last abdominal segment is constricted. This character is listed as a synapomorphy for the Eriopterinae but apparently does not belong to the groundplan condition of Hexatominae. 77. Respiratory horn club-shaped 78. Mandible single toothed in the middle Conosia and Eutonia are considered sistergroups on the basis of their club-shaped respiratory horns (figs. 86, 92) and single toothed mandibles (figs. 66-68). The latter character occurs also in some species of Hexatoma (Alexander 1920). 79. Larval creeping welts Dorsal and ventral creeping welts on segments 5-10 are found in larvae of Eloeophila, Limnophila (Lasiomastix) macrocera, Conosia, Eutonia and ap- parently also Neolimnomyia (Brachylimnophila) (see Beling 1886). 80. Markings on ventral spiracular lobes divided into two This character is present in Paradelphomyia and a number of higher Hexatominae including Neo- limnomyia (Brachylimnophila) (figs. 57, 72, 78). It does not occur in all species of Limnophila and is unknown for Neolimnomyia (Neolimnomyia) where the spiracular disc seems to be permanently closed (fig. 84; Brindle & Bryce 1960). 81. Pupal abdominal segments with lateral carina Distinct lateral carinae provided with spines (fig. 89) are present on the abdominal segments of the pupae of Pilaria, Phylidorea, Euphylidorea, Eloeophila, Neolimnomyia and several species of Limnophila. 82. Penultimate segment of larvae inflated Among Hexatominae inflation of the penulti- mate segment seems to be weak or absent in a number of genera. Distinct inflation is known to occur in Paradelphomyia, Pilaria, Phylidorea, Eu- phylidorea, Eloeophila (fig. 85), Hexatoma (fig. 64) and Limnophila. 242 7. PEDICIINAE Introduction Pediciinae is a small subfamily, divided in two tribes, Ulini and Pediciini. The larvae of Ula (Ulini) feed on fungi. The larvae of the other genera discussed here belong to the Pediciini and are predators on oligochaetes, mites, insect larvae and the like (details in Crisp & Lloyd 1954 and Lindner 1959). Because of the very massive head capsule, Pediciini are not able to swallow large prey as a whole but have to tear them to pieces (Lindner 1959). Pupae of Pediciinae are enclosed in a sub- stantial silken tube covered with detritus, as in Limoniinae (character 9). The adopted phylogeny (fig. 7) shows two groups of Tricyphona. The palaearctic species T. immaculata, T. schummeli and T. unicolor are in- cluded in Tricyphona 1; T. inconstans from the Nearctic represents Tricyphona 2. Rhaphidolab- ma, generally placed as a subgenus of Dicranota (Savchenko et al. 1992), is placed as sistergroup of Tricyphona 2. Characters 83. Cardo reduced In Pediciini the cardo is reduced to a small plate (fig. 100). The reduced inner lobe of the maxilla is closely aligned to the outer lobe. According to Crisp & Lloyd (1954) reduction of the inner lobe is less in Pedicia rivosa, intermediate in Tricyphona spec., and most obvious in Dicranota bimaculata. The available information on the other species and genera does show that reduction of the inner lobe is rather variable and apparently not linked to the adopted phylogeny. 84. Larvae with ventral creeping welts on sternites 6-10 or 7-10 Ventral creeping welts on the anterior part of sternites 5-10 is interpreted as the plesiomorphic condition in Tipuloidea (character 29). In Pediciini welts are present on sternites 7-10 (figs. 107, 108), or as pseudopods in Raphidolabis, Paradicranota and Dicranota on sternites 6-10 (fig. 104). On the basis of characters 88 to 94 it is assumed that the number of welts decreased from six to four in Pediciini with an increase to five in the Raphido- labis-Dicranota lineage. Of interest is the presence of welts on segments 5-10 in the apparently prim- itive pediciine genus Sigmatomera (see chapter 9). 85. Hypostomal plates with three or four teeth 86. Two parts of hypostoma with a push-button connection The hypostoma of Pediciini is very characteristic (fig. 97). It is divided along the midline, each half with three or four teeth and the two parts are held together with a push-button connection. Usually one or two buttons are present, in Tricyphona 2 there are three. A similar connection is present in Pedicia, but without a clear separation in push- buttons. 87. Spiracular field with prominent ventral lobes. 88. Ventral spiracular lobes very elongate In Pediciini only the ventral spiracular lobes are prominent (figs. 105-108), and very elongate ex- cept for Tricyphona 1 (figs. 105, 107). The ventral lobes terminate with a group of setae at the end. 89. Mandible sickle shaped As in higher Hexatominae (character 66), the mandibles of Pediciini are sickle shaped (fig. 102), Tricyphona 1 excepted (fig. 101). They are pro- vided with four large ventral teeth and the lower ventral tooth is always the largest and more or less rectangular. 90. Spiracular field small and spiracles close to- gether The spiracular field in Tricyphona 1 is large with the spiracles far apart (fig. 105), as in Ula and the other genera of the Dactylolabis-Tipulidae lineage. In the remaining Pediciini the spiracular field is small and the spiracles are close together (fig. 106). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 93-94. Head capsule of Tricy- phona immaculata. - 93, dorsal view; 94, ventral view (Lindner 1959) (fig- ures redrawn). 91. Creeping welts concave in the middle The creeping welts of Tricyphona 1 are as found in the other Tipuloidea. In the remaining Pediciini they are laterally elevated into humps or pseudop- ods (fig. 104; character 95). 92. Pupal pleura with circular areas of spicules The abdominal pleura of the pupae of Tricy- phona 2 and Rhaphidolabina are provided with large, circular areas of short rows of microscopic spicules. It is not known whether this character is present in Tricyphona 1. 93. Hypostomal plates with four teeth Each hypostomal plate in Tricyphona 2 and Rha- phidolabina is provided with four teeth, whereas the other Pediciini possess three teeth. 94. Spiracles on elevations Pedicia, Rhaphidolabis, Paradicranota and Di- cranota larvae have the spiracles situated on small elevations (fig. 108), a character not found among other Pediciinae or Tipuloidea. The small eleva- tions are interpreted by Crisp & Lloyd (1954) as representing the lateral spiracular lobes. 95. Pseudopods Pseudopods, apically set with rings of recurved hooklets (fig. 104) are found in the larvae of Rha- phidolabis, Paradicranota and Dicranota. They are 243 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 95-108. Pediciini, larval characters. - 95, Tricyphona immaculata, head capsule of first instar, dorsal view; 96, idem, ventral view; 97, T. immaculata, hypostoma (ap. apophyse), ventral view; 98, Dicranota (Rhaphidolabina) flaveola, antenna; 99. Dicranota (Dicranota) bimaculata, antenna; 100, T. immaculata, right maxilla (c, cardo), ventral view; 101, idem, left mandible, dorsal view; 102 Pedicia rivosa, left mandible, dorsal view; 103, T. immaculata, prementum, ventral view; 104, Dicranota (Paradicranota) subtilis, creeping welts (pseudopods) of segments 8 and 9; 105, T. immaculata, terminal segment, dorsal view; 106, Dicranota spec., terminal segment, dorsal view; 107, T. immaculata, larva, lateral view; 108, Pedicia spec., larva, lateral view (fig. 95, 96: Okely 1979; 97, 100, 101, 103-105, 107: Lindner 1959; 98: Alexander 1920; 99: Reusch 1988; 102: Wardle & Taylor 1926; 106: Savchenko 1986; 108: Alexander & Byers 1981) (all figures redrawn). present on the anterior part of sternites 6-10. The only other genus of Tipuloidea with pseudopods is Aphrophila (see chapter 9). 96. Pupae with ventral creeping welts Pupae of Rhaphidolabis are not known. Those of the other taxa possess dorsal creeping welts, situ- ated more lateral in Paradicranota and Dicranota. Ventral creeping welts are present in Tricyphona and Pedicia, whereas in Paradicranota and Dicra- nota the sternites are provided with fleshy projec- 244 tions, corresponding with the larval pseudopods and used to assist the movements of the pupa through the mud (Miall 1893). 97. Apical papillae of antenna reduced Pediciinae usually have the larval antenna pro- vided with two distinct apical papillae (figs. 98, 114; character 7). In Rhaphidolabis and Dicranota the papillae are short (fig. 99; Alexander 1920, Reusch 1988) and in Tricyphona immaculata only one short papilla is present (information for the other species assigned to Trichypona | is not avail- able). 98. Basal segment of antenna short The basal antennal segment of Tipuloidea larvae is usually well developed, but short in Uda (fig. 114) and some species of Dactylolabis (see character 7). In Pediciini the basal segment is elongate (fig. 99) except for Tricyphona 2 and Rhaphidolabina (fig. 98) with a length intermediate between Ula and the other Pediciini. 8. LIMONIINAE Introduction Palaearctic genera of Limoniinae are divided into two tribes, Antochini and Limoniini (Dienske 1987, Savchenko et al. 1992). This division is also apparent on the basis of immature characters. In figure 8 the genera Atypophthalmus to Discobola belong to the Limoniini, the others to the Antochini. The phylogenetic arrangements of the genera dealt with is based on few characters and distinct sistergroup relations within the Limoniini could not be established. This is at least of some interest because genera of Limoniini are usually considered subgenera of Limonia by most North American and British workers. It appears that, as far as char- acter polarities could be established within the Li- moniini, all character-states are present in repre- sentatives of the large genus Dicranomyia. Limoniinae larvae are herbivorous, feeding on decayed plant material, diatoms, organic mud and the like. Larvae of several species live in fungi (Lindner 1958). Experiments to find out their pref- erences for leaf-litter of different deciduous trees are described by Lindner (1959). Characters 99. Larvae in tube or case Unlike other Tipuloidea, larvae of Limoniinae live inside silken or gelatinous tubes, usually co- vered with detritus, silt or diatoms, some species of Dicranomyia excepted (Wood 1952: D. peringueyi, Brindle 1967: D. didyma). Thaumastoptera larvae live in a dark grey or blackish hardened elliptical flattened case. Information is not available for Aty- pophthalmus. 100. Pupal creeping welts elliptical In the Limoniinae the pupal creeping welts are distinctly elliptical, a character not found in other Tipuloidea. The central part of the welts is fre- quently naked or bears a reduced number of spic- ules (figs. 157-159). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea 101. Spiracles oblong or elliptical 102. Spiracles placed obliquely Antocha larva are apneustic, without spiracles. The spiracles of the other Limoniinae are oblong or elliptical and placed obliquely (figs. 141, 142, 150). These two characters must be considered weak synapomorphies because they are found, al- beit less pronounced, in several other Tipuloidea (figs. 78, 105, 136). 103. Larval segments 2 to 10 with creeping welts In addition to the creeping welts present on segments 5-10, the genera included in the Limo- niini and Elliptera possess creeping welts, but with a lower relief, on the anterior margin of segments 2,3 and 4 and the posterior margin of segment 10 (figs. 143, 144). It is not known whether this char- acter is present in Atypophthalmus. 104. Spiracular lobes reduced Limoniini have the spiracular lobes reduced and very frequently they are indicated only by the dark patches of chitinization or by the general outline of the spiracular field (figs. 141-144). 105. Caudal end of head capsule tridentate Only in Limoniini the caudal end of the head capsule is tridentate (figs. 139, 140). The space between the lateral and central points is inter- preted by Lindner (1959) as remnants of the fron- tal sutures. This character remained unknown for Atypophthalmus and is not present in all species of Limonia. 9. REMAINING GENERA Discussed below are a number of genera which could not be incorporated among the genera men- tioned in figs. 4-8 because of the limited, or in some cases controversial, information available. The cur- rent systematic position of the taxa is given in parentheses. Aphrophila (Eriopterinae). - Winterbourn & Gregson (1989) figure the larva of Aphrophila neozelandica, which is very aberrant compared to the known larvae of Eriopterinae. The last abdom- inal segment appears constricted (character 36) and the head capsule is dissected (character 49). Spiracular lobes are absent (character 41) and the larva seems to be apneustic. Unlike other Eriopte- rinae, dorsal creeping welts (‘scars’) are present on tergites 6-10. Sternites 5-10 also possess creeping welts (character 48), but shaped as pseudopods very similar to the pediciine taxa Rhaphidolabis, Paradicranota and Dicranota (character 95). 245 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 109-118. Ula, larval characters. - 109, U. mollissima, larva, lateral view; 110, idem, head capsule, dorsal view; 111, idem, ventral view; 112, U. sylvatica, head capsule of first instar larva, dorsal view; 113, U. mollissima, left maxilla (c, cardo), ventral view; 114, U. elegans, antenna; 115, U. mollissima, left mandible, dorsal view; 116, U. sylvatica, head capsule, dorsal view; 117, idem, spiracular disc of first instar larva; 118, U. mollissima, spiracular disc (figs. 109- 113, 115, 117, 118: Lindner 1959; 114: Alexander 1920; 116: Bryce 1957b) (all figures redrawn). Gonempeda (Eriopterinae: Eriopterini). - The only original figures of the pupa of Gonempeda flava are those by Brindle (1967, as Cheilotrichia) and only a few characters can be interpreted. The respiratory horns are not earshaped and the me- sonotal crest seems to be lacking (characters 38, 46 and 50 not present). Hesperoconopa (Eriopterinae: Molophilini). - The larva and pupa of Hesperoconopa dolicho- phallus are described and figured by Hynes (1968). The unique terminal segment of the larva is setu- lose, elongate cone shaped and bifurcate at the extreme tip, lacking spiracles. Information is avail- 246 able for the following characters: Present are 36, 38, 41 (but abdominal end quite different from Cladura and Chionea), 49 and 59 (ventral bars not apically toothed). Not present are 42 (palpsheaths appear upcurved), 48, 50 and 53. Neolimnophila (Eriopterinae: Cladurini). - Bel- ing (1886) presents a description of the larva and pupa of Neolimnophila carteri (as Limnophila byalipennis). The pupal respiratory horns are ab- sent (character 38) and the rounded spiracular field is without lobes (character 41). This, together with the absence of characters 49 and 50 supports the systematic position of Neolimnophila among the OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea as my 07 MIN N 124 My 07 MN 4777) 123 Zz 2 MA zi £ A ND Dit 1 ES 6/ Ze 3 ; | À Ò Figs. 119-126. Austrolimnophila, larval characters. - 119, A. ochracea, head capsule (cs, coronal suture; fs, frontal suture), dorsal view; 120, idem, ventral view; 121, A. medialis, left maxilla (mp, maxillary palp), ventral view; 122, A. ochracea, hypopharynx (hp), hypopharyngeal bar (hb) and prementum (pm); 123, idem, spiracular disc; 124, idem, hypostoma; 125, idem, right mandible, dorsal view; 126, idem, labrum, dorsal view. - Figs. 127-128. Dactylolabis, larval characters. - 127, D. sexmaculata, spiracular disc; 128, D. transversa, left mandible, dorsal view. — Figs. 129-130. Epiphragma, larval characters. - 129 E. ocellare, labrum, dorsal view; 130, E. fasciapenne, terminal segment, oblique posterior view (figs. 119, 120, 122-125: Lindner 1959; 121, 126: Wood 1952; 127-129: Brindle & Bryce 1960; 130: Alexander & Byers 1981) (all figures redrawn). 247 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 131 132 | A NS 135 i i a 138 Figs. 131-132. Dicranoptycha pallida, larval characters. - 131, head capsule, ventral view; 132, hypopharynx and hypopharyngeal bar. - Figs. 133-137. Helius longirostris, larval characters. - 133, right maxilla, ventral view; 134, head capsule, dorsal view; 135, hypostoma (ap, apophyse); 136, spiracular disc; 137, larva, lateral view. - Fig. 138, Limnophilomyia lacteitarsus, left mandible, dorsal view (figs. 131, 132: Alexander 1919b; 133-137: Cramer 1968; 138: Wood 1952) (all figures redrawn). Cladurini, although character 53 seems to be pres- ent. Sigmatomera (Eriopterinae). - Hudson (1920) provides information for Sigmatomera (Austrolimnobia) rufa. Larvae are living in the semi-liquid vegetable detritus at the stem of the epiphytic Astelia leaves and the pupa is enclosed in an upright, rather tough, extremely elongate silken tube. Alexander (1930) adds interesting notes on the biology of the larva of Sigmatomera (Sigma- 248 tomera) shannoniana. The larva is predacious and lives in rot holes of trees where it makes perman- ent slimy galleries in the debris, cementing the particles together with the same slimy substance. The prey are Culicidae larvae, which are tremend- ously eager to feed on the slimy substance and, while feeding, are devoured by the Sigmatomera larva. The combination of predatory larva and pupa in a silken tube is found only in Pediciini. The larva of rufa has dorsal and ventral creeping welts on segments 5-10 as in Ula and several other Tipuloi- dea, whereas Pediciini have a reduced number of creeping welts (character 84), indicating that Szg- matomera represents one of the most basal line- ages of the Pediciini. Teucholabis (Eriopterinae: Cladurini). - In Teu- cholabis complexa, immature stages of which are described and figured by Alexander (1920), the following characters are present: 37 (fig. 52), 42, 49, 50, 53. Not present are: 36, 38 (the respiratory horns are short), 43 and 56. Information about the other characters is not available. Teucholabis is usually considered to belong to the Cladurini but the characters of the larva and pupa point in the direction of Eriopterini or Molophilini, in spite of the aberrant three lobed spiracular field. Trentepohlia (Eriopterinae: Gonomyiini). - The limited amount of information available for Tren- tepohlia is reviewed by Alexander (1920) and points towards a relationships with Limnophilo- myia. Both genera are usually considered to belong to the Eriopterinae but in fig. 4, Limnophilomyia is placed next to the Limoniinae, at about the po- sition suggested by Rogers & Byers (1956). Char- acters in common in Trentepohlia and Limnopht- lomyia are reduction of the spiracular lobes, very elongate anal papillae divided by constrictions (as in several species of Lipsothrix (Hynes 1965) and Hexatominae, see character 72), inner leg sheaths of the pupa the longest (character 33). Elephantomyia. - This genus has been placed with a wide array of genera and tribes: in Limnob- ina anomala (Osten Sacken 1869), in Antochini (Alexander 1919a), with Toxorrhina in the Ele- phantomyiini as part of the Eriopterinae (Alex- ander 1920), in Elephantomyiini as part of the Hexatominae (Alexander 1927; Savchenko 1983a), in Limnophilini as part of the Hexatominae (Sav- chenko & Krivolutskaja 1976), in Heliini or Ele- phantomyiini as part of the Limoniinae (Lacksche- witz 1932; Lackschewitz & Pagast 1940-1942; Savchenko et al. 1992). Immature stages of five Elephantomyia species are known. Larvae and pupae are described and figured by Alexander (1920) for the type-species westwoodi, by Bangerter (1934) for edwardsi and by Wood (1952) for montana and aurantiaca, with the additional note that the pupa of pseudosimilis closely resembles that of montana. Elephantomyia is the only genus of Tipuloidea with two strikingly different types of larvae. These differences are coupled with the larval habitat. The species aurantiaca and pseudosimilis live in moss cushions in waterfalls or along the edges of streams OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea and tricklets. The larva of pseudosimilis is un- known, but the larva of aurantiaca has a massive head capsule of ‘Limonian construction’ (Wood 1952). The larvae of the other three species live in decaying wood. Their head capsule is minute, nar- row, consisting of three elongate plates, and very different from the 'Limonian type’, or any other type of head capsule. The amount of difference is so large that one is inclined to believe that at least different genera are involved. However, Elephan- tomyia is most probably monophyletic on account of characters of the adults. Furthermore, the four described larva are the only Tipuloidea with creep- ing welts restricted to sternites 8 to 10. The three species living in decaying wood (westwoodi, edwardsi and montana), share the fol- lowing characters: (A.) Body tapering toward either end but more abruptly anteriorly with the head capsule protruding as a sharp narrow point. - (B.) Sternites 8-10 anteriorly with a very conspic- uous mouth-like depression. — (C.) Sternite 11 with a transverse mouth, lying near end of seg- ment, its anterior lip with a dense fringe of long hairs directed backward. According to Bangerter (1934) the four delicate anal papillae can be re- tracted into the opening, to be closed off by the long hairs. - (D.) Spiracular disc surrounded by four lobes (fig. 172). - (E.) Head capsule (fig. 171) very small, narrow and elongate. - (F.) Mandibles very small (see for details Bangerter 1934). - (G.) Men- tal region feebly chitinized. - (H.) Oesophageal region strengthened with oblique parallel ridges of chitin, producing a latticed effect. - (I.) Antennal papilla large, suboval. - (J.) Sides of head capsule on swelling behind the maxilla with a brush of very long hairs (not certain for montana). - (K) Pupal legsheaths with fore femora distinctly swollen. This last character is mentioned for montana by Wood (1952: 247) and it must be assumed that it also present in pseudosimilis. The species west- woodi and edwardsi have the larval thoracic and abdominal segments with a dorsal and ventral transverse ridge of short, stiff hairs just before the caudal margins and a conspicuously elongate bristle on the ventral spiracular lobes (fig. 172). Of the above mentioned characters, characters D and G are found in other Tipuloidea as well. All other characters are unique and of little help in establishing the phylogenetic position of Elephan- tomyia. On the other hand, all five species have the pupal abdominal segments unarmed (character 14), and aurantiaca possesses a massive head cap- sule with a complete, five-toothed hypostoma (fig. 170, character 11), a toothed prementum (charac- ter 16; ventral plate of hypopharynx in Wood 1952) and a reduced spiracular field as in Limoniini (fig. 169, character 104). These characters indicate 249 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 m Im S Figs 139-142. Limoniinae, larval characters. - 139, Limonia macrostigma, head capsule, ventral view; 140, idem, dorsal view; 141, Dicranomyia modesta, spiracular disc; 142, L. macrostigma, spiracular disc (Lindner 1959) (figures redrawn). that at least aurantiaca, but probably Elephanto- myta as a whole, belongs to the Limoniinae. Ac- cording to Wood (1952), the aurantiaca larva has a head capsule resembling that of Geranomyia sex- ocellata, from which it differs only in the armature of the mentum and the antenna. Atarba (Hexatominae). - Larva and pupa of Atarba picticornis are described by Rogers (1927a), who stated that ‘on the basis of larva and pupa characters, Atarba very clearly is among the genera whose relationships are with the Limoniini [=Limoniinae]’. On adult characters, Atarba is generally considered a tribe of its own, placed in the 250 Hexatominae (Byers 1981, Savchenko et al. 1992). Atarba picticornis possesses a complete hypos- toma, as found in Ula and the Dactylolabis-Tipu- lidae lineage only. Of the characters mentioned in figure 4, no information is available for characters 8, 10, 12, 18, 19 and 35. Of the other characters, 4, 5, 14, 15, 16, probably 17 (interno-lateralia partly fused), 26, 28, 29 and 30 (but segments 3-11 with a unique arrangement of very low creeping welts) and 34 are present. Apart from character 26 (pe- nultimate segment inflated), this strongly indicates that Atarba might indeed represent one of the most basal lineages of the Limoniinae, in figure 8 branching of between characters 9 and 99. Charac- OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Figs. 143-152. Limoniinae, larval characters. - 143, Dicranomyia modesta, larva, lateral view; 144, Limonta macros- tigma, larva, lateral view; 145, idem, labrum and antennae, dorsal view; 146, idem, hypostoma; 147, Dicranomyia trifilamentosa, hy popharyngeal teeth (hp) and prementum (pm), ventral view; 148, Metalimnobia bifasciata, premen- tum, rudimentary labial palps (Ip) and part of hypopharynx (displaced laterally), ventral view; 149, Dicranomyia monostromia, cross section through anterior part of larval head, showing labrum (lb), epipharynx (ep), hypopharynx (hp), prementum (pm) and hypostoma (hs); 150, Orimarga mirabilis, spiracular disc; 151, Dicranomyia autumnalis, left maxilla (sp, separate sclerotized plate on inner lobe), ventral view; 152, D. modesta, left maxilla (sp, separate sclerotized plate on inner lobe), ventral view (figs. 143-146, 152: Lindner 1959; 147: Tokunaga 1933: 148, 151: Cramer 1968; 149: Tokunaga 1930; 150: Rogers 1927c) (all figures redrawn). 2511 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 153-159. Pupae. - 153, Limnophilomyia lacteitarsus, female, lateral view; 154, Lipsothrix sylvia, female, ventral view; 155, Austrolimnophila medialis, female, lateral view; 156, Phalacrocera replicata, female, laterodorsal view; 157, Metalimnobia bifasciata, female, lateral view; 158, Dicranomyia trifilamentosa, female, lateral view; 159, Antocha saxicola, female, lateral view (figs. 153, 155: Wood 1952; 154: Rogers & Byers 1956; 156: Peus 1952; 157: Cramer 1968; 158: Tokunaga 1933; 159: Alexander 1920) (all figures redrawn). ters listed exclusively for the Eriopterinae (fig. 5), Hexatominae (fig. 6), Pediciinae (fig. 7) and Limo- niinae (fig. 8) are not present except for character 37 (pupa with middle leg sheaths the shortest). Idioptera, Rhamphophila and Tonnoiraptera (Hexatominae). - The information provided for Idioptera pulchella (by Brindle & Bryce 1960, Brin- dle 1967), Rhamphophila obscuripennis (by Hud- son 1920, as Limnophila sinistra) and Tonnoirap- tera neozelandica (by Tonnoir 1926, as Alexandrella) is very limited. According to Brindle & Bryce the larva of I. pulchella closely resembles the larva of Phylidorea. Typical hexatomine char- acters are not mentioned for R. obscuripennis but seem to be present in T. neozelandica (Tonnoir 1926: ‘the head armature is typically that of the tribe Hexatomini [=Hexatominae] of the carni- 252 vorous group, with elongated and very much dis- sected capsule and with sharp, curved mandibles’). Achyrolimonia and Neolimonia (Limoniinae: Limoniini). — The information on the larva of Neo- limonia dumetorum (Beling 1886) and Achyroli- monia decemmaculata (Brindle 1967) shows that characters 101, 102 (spiracles oblong and placed obliquely) and 104 (spiracular lobes reduced) are present. Therefore, both genera are to be placed among the Limoniini. 10. INDEX TO THE LITERATURE Information from the literature is found in a wide variety of publications and in various forms (notes, descriptions, keys, figures). A number of the more important publications are mentioned in OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea 17/0) za 172 4 DN Figs. 160-168. Cylindrotominae, larval characters. - 160, Phalacrocera replicata, head capsule (c, cardo; ps, premax- illary suture), ventral view; 161, idem, left mandible of first instar larva, ventral view; 162, idem, right mandible of last instar larva, from inside; 163, idem, left maxilla (c, cardo; sp, separate sclerotized plate on inner lobe), oblique ventral view; 164, Diogma glabrata, spiracular field; 165, P. replicata, hypostoma (hs) of first instar larva, ventral view; 166, idem, hypostoma (hs) of full grown larva, ventral view; 167, 168, idem, hypopharynx (hp) and prementum (pm), oblique frontal and lateral view. - figs. 169-172. Elephantomyia, larval characters: 169, E. awrantiaca, abdominal end, lateral view; 170, idem, hypostoma; 171, E. edwardsi, anterior part of head capsule, ventral view; 172, E. westwoodi, spiracular field (figs. 160, 163, 164, 167, 168: Peus 1952; 161, 162, 165, 166: Bengtsson 1897; 169, 170: Wood 1952; 171: Bangerter 1934; 172: Alexander 1920) (all figures redrawn). 253 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 173-180. Tipulidae, larval characters. - 173, Brachypremna dispellens, head capsule (c, cardo; ps, premaxillary suture), ventral view; 174, Holorusia spec. (from Brazil, cf. j#juyensis), epipharynx (tm, torma; eb, epipharyngeal bar), lateroventral view; 175, 176, Tipula livida, hypopharynx (hp) and prementum (pm), ventral and lateral view; 177, Prionocera turcica, spracular disc; 178, T livida, right maxilla (c, cardo; sp, separate sclerotized plate on inner lobe), ventral view; 179, 180, Nephrotoma spec., left mandible, ventral and dorsal view (fig. 173: Gelhaus & Young 1991; 174: original; 175, 176, 178: Chiswell 1955; 177: Theowald 1957; 179, 180: Savchenko 1983b) (all figures redrawn). chapter 2. Not all of these publications are indexed all the available literature containing relevant in- here because some of them are compilations only. formation. The authors are grateful for additions to Genera, subgenera and species are listed alphabet- this index. In the index the following marks are ically. An effort has been made to give reference to used: 254 + Type-species. V! Preserved material available, Zoological Mu- seum, Amsterdam. Larva. Pupa. Identification of pre-imaginal stages not cer- tain. 29 Ino} |e! Achyrolimonia decemmaculata Loew: Brindle 1967 (LP), Savchenko 1985 (LP). Afrolimonia: see under Libnotes. Amalopis: see under Pedicia. Antocha (Antocha) bifida Alexander: Hinton 1966, 1968 (P). monticola Alexander: Needham & Christensen 1927 (LP) + saxicola Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (LP), Peterson 1960 (L), V! (L). vitripennis Meigen: Hinton 1957 (P), Brindle 1967 (LP), Hinton 1968 (P), Rozkosny & Pokorny 1980 (L), Savchenko 1985 (LP), V! (LP). spec.: Malloch 1917 (L) (as Genus incertus 2), Vimmer 1928 (LP, as calceata), Alexander 1931 (LP), Byers 1978 (L), Alexander & Byers 1981 (L), Savchenko 1986 (L). Antocha (Orimargula) + alpigena Mik: Bangerter 1929 (LP), Rozkosny & Po- korny 1980 (L), Savchenko 1985 (LP). australiensis Alexander: Hinton 1965, 1968 (P). Aphrophila neozelandica Edwards: Winterbourn & Gregson 1989 (L). Arctoconopa carbonipes Alexander: Hynes 1969a (LP), Byers 1978 (L), Savchenko 1982 (LP). Atarba + picticornis Osten Sacken: Rogers 1927a (LP). viridicolor Alexander: Rogers 1927a (P). spec.: Byers 1981 (L). Atypophthalmus inustus Meigen: Beling 1878 (LP, as obscuricornis), V! (P). + umbratus de Meijere: de Meijere 1917 (L). Austrolimnobia: see under Sigmatomera. Austrolimnophila medialis Alexander: Wood 1952 (LP). ochracea Meigen: Beling 1873a (LP, as pallida), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP), V! (P). Baeoura claripennis Alexander: Wood 1952 (LP). witzenbergi Wood: Wood 1952 (P). Brachylimnophila: see under Neolimnomyia. Cheilotrichia (Empeda) cinerascens Meigen: *Levy 1919 (LP), Crisp & Lloyd 1954 (L), Lindner 1959 (L), Brindle 1967 (L), Krivo- sheina 1969 (L), Savchenko 1982 (P), 1986 (L), Reusch 1988 (LP). Chionea alexandriana Garrett: Byers 1983 (P). + araneoides Dalman: Egger, Frauenfeld & Brauer 1854 (L), Alexander 1920 (L), Savchenko 1982 (L). lutescens Lundström: Edwards 1936 (P), Savchenko 1982 (P), Byers 1983 (P). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea scita Walker: Byers 1983 (L). stoneana Alexander: Byers 1983 (L). Cladura + flavoferruginea Osten Sacken: Alexander 1920, 1922 (LP). Conosia + irrorata Wiedemann: Wood 1952 (LP). Crunobia: see under Pedicia. Cryptolabis magnistyla Alexander: Hynes 1963 (LP). spec.: Byers 1978 (L), Alexander & Byers 1981 (L). Cylindrotoma + distinctissima distinctissima Meigen: Lenz 1920b (LP), Haake, 1922 (L), Peus 1952 (LP), Brindle 1967 (LP), Brodo 1967 (LP). distinctissima americana Coquillett: Cameron 1918 (LP, as splendens), Alexander 1920 (LP, as splendens). Dactylolabis cubitalis Osten Sacken: Alexander 1920 (P), Johannsen 1934 (P). denticulata Bergroth: Mik 1894 (LP), Alexander 1920 (LP), Vimmer 1925 (LP), Bangerter 1931 (LP), Jo- hannsen 1934 (L), Wesenberg-Lund 1943 (L). hudsonica Alexander: Sinclair 1988 (LP). montana Osten Sacken: Sinclair 1988 (LP), V! (LP). sexmaculata Macquart: Bangerter 1931 (LP), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP). transversa Meigen: Bangerter 1931 (LP, as gracilipes), Brindle & Bryce 1960 (L), Brindle 1967 (L), Savchenko 1986 (LP). wodzickii Nowicki: Nowicki 1867 (LP), Alexander 1920 (LP), Lindner 1959 (L). spec.: Alexander & Byers 1981 (key). Dicranomyia (Dicranomyia) autumnalis Staeger: Cramer 1968 (LP), Savchenko 1985 (LP), 1986 (LP). capicola Alexander: Wood 1952 (LP). casei Alexander: Williams 1943 (LP). chorea Meigen: Reusch 1988 (L), V! (P). didyma Meigen: Engel 1916 (LP, as trinotata), Vimmer 1925 (LP, as trinotata), Ussing 1929 (LP), Bryce 1957a (L), Brindle 1967 (LP), Savchenko 1985 (LP), V! (P). floridana Osten Sacken: Rogers 1932 (LP). frontalis Staeger: Reusch 1988 (LP). grimshawi Alexander: Williams 1943 (LP). halobia Tokunaga: Tokunaga 1936 (P). humidicola Osten Sacken: Alexander 1920 (LP, as ba- dia), Johannsen 1934 (P, as badia), Byers 1978 (L). jacobus Alexander: Williams 1943 (LP). kauaiensis Grimshaw: Swezey 1915 (LP, as foliocunic- ulator). mariana Seligo: Seligo 1931 (LP). marmorata Osten Sacken: Saunders 1928 (LP, as sig- nipennis). mitis Meigen: Vaillant 1956 (L), Brindle 1967 (L), Savchenko 1985 (L). + modesta Meigen: Vimmer 1925 (LP), Lindner 1959 (L), Brindle 1967 (LP), Savchenko 1985, 1986 (L). monostromia Tokunaga: Tokunaga 1930 (LP), Kaw- ada 1960 (LP), Hinton 1968 (P). nigrescens Hutton: Winterbourn & Gregson 1989 (L). peringueyi Alexander: Wood 1952 (LP). punctulata De Meijere: Alexander 1931 (LP). sera Walker: Szadziewski 1979 (LP). stulta Osten Sacken: Alexander 1920 (LP), Johannsen 255 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 1934 (L). tipulipes Karsch: Wood 1952 (LP). trifilamentosa Alexander: Tokunaga 1933 (LP), Kaw- ada 1960 (LP), Hinton 1968 (P). ventralis Schummel: Brindle 1967 (LP), Savchenko 1985 (LP). vicina Macquart: Hemmingsen et al. 1959 (L, as cana- riensis) Dicranomyia (Idioglochina) + marmorata Osten Sacken: Hinton 1968 (P), Ring 1978 (P). tokunagana Alexander: Tokunaga 1939 (L, as glori- 054). Dicranomyia (Idiopyga) melleicauda complicata De Meijere: V! (P). Dicranophragma: see under Limnophila. Dicranoptycha megaphallus Alexander: Young 1987 (LP) minima Alexander: Alexander 1919b (L). pallida Alexander: Alexander 1919b (LP, as winne- mana). spec. Alexander 1920 (LP). Dicranota (Dicranota) bimaculata Schummel: Miall 1893 (LP), Malloch 1917 (LP), Levy 1919 (LP), Vimmer 1925 (LP), Johannsen 1934 (L), Wesenberg-Lund 1943 (L), Crisp & Lloyd 1954 (L), Brindle 1967 (LP), Reusch 1988 (LP), V! (L). + guerini Zetterstedt: *Fahy 1972 (L). spec.: Vimmer 1906 (L), 1925 (P), Malloch 1917 (L), Vimmer 1924 (L, as Amalopis), Miall 1934 (LP), Cook 1949 (L), Savchenko 1986 (L), Reusch 1988 (L), V! (L). Dicranota (Paradicranota) robusta Lundstròm: Brindle 1962, 1967 (LP), Sav- chenko 1986 (LP). subtilis Loew: Lindner 1959 (L), Brindle 1962, 1967 (L), Savchenko 1986 (L). Dicranota (Rhaphidolabina) claripennis Verrall: Brindle 1962, 1967 (L). + flaveola Osten Sacken: Alexander 1920 (LP). Dicranota (Rhaphidolabis) cayuga Alexander: Alexander 1920 (L). exclusa Walker: Nielsen et al. 1954 (L), Brindle 1962 (L). + tenuipes Osten Sacken: Needham 1908 (L), Alexander 1920 (L). Diogma + glabrata Meigen: Müggenberg 1901 (LP), Alexander 1920 (LP), Lenz 1920b (LP), Haake 1922 (LP), Peus 1952 (LP), Brindle 1967 (LP), Krivosheina 1969 (L). spec.: Malloch 1917 (L), Byers 1978 (L). Diotrepha: see under Orimarga. Discobola + annulata Linnaeus: Krivosheina 1969 (L), Savchenko 1985, 1986 (L), Krivosheina et al. 1986 (L). caesaria Osten Sacken: Mik 1884 (P), Alexander 1920 (P). Elephantomyia aurantiaca Alexander: Wood 1952 (LP). edwardsi Lackschewitz: Bangerter 1934 (LP), Sav- chenko 1986 (LP). montana Alexander: Wood 1952 (LP). pseudosimilis Alexander: Wood 1952 (P). + westwoodi Osten Sacken: Alexander 1920 (LP). spec.: Malloch 1917 (L) (as Genus incertus 1), Krivo- sheina 1969 (L), Savchenko 1986 (L). Elliptera + omissa Schiner: Mik 1886 (LP), Malloch 1917 (LP), Alexander 1920 (LP), Vimmer 1925 (L), Bangerter 1934 (LP), Johannsen 1934 (LP), Brindle 1967 (LP), Savchenko 1985 (LP), V! (LP). Ellipteroides (Ellipteroides) lateralis Macquart: Bangerter 1928 (LP), Brindle 1967 (LP), Savchenko 1982 (LP). Ellipteroides (Progonomyia) nigrobimbo Alexander: Wood 1952 (LP). Ellipteroides (Protogonomyia) alboscutellatus von Roser: Bangerter 1930 (LP), Brin- dle 1967 (LP), Savchenko 1982 (LP). Eloeophila (= Ephelia) apicata Loew: Brindle & Bryce 1960 (L), Brindle 1967 (LP). dubiosa Alexander: Wood 1952 (LP). + maculata Meigen (= marmorata Meigen): Wolff 1922 (L), Bangerter 1928 (LP), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Pokorny 1978 (LP), Savchenko 1986 (LP), Reusch 1988 (P), V! (L). mundata Loew: Brindle & Bryce 1960 (L), Pokorny 1978 (LP), Savchenko 1986 (LP). submarmorata Verrall: Crisp & Lloyd 1954 (L), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Pokorny 1978 (LP), Savchenko 1986 (LP). trimaculata Zetterstedt: Brindle & Bryce 1960 (L), Brindle 1967 (LP). verralli Bergroth: Brindle 1958 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Reusch 1988 (P). spec.: Walton 1944 (L, as undescribed, erroneously considered Pedicia rivosa by Hinton 1950), Alexander & Byers 1981 (L, as Limnophila spec.), V! (L). Elporiomyia: see under Limnophila. Empeda: see under Cheilotrichia. Epiphragma + fasciapenne Say: Needham 1903, 1908 (LP), Malloch 1917 (LP), Alexander 1920 (LP), Teskey 1976 (L), Alexander & Byers 1981 (L). imitans Alexander: Bruch 1939 (LP). ocellare Linnaeus: Beling 1873b (LP, as picta), *Wolff 1922 (L, as punctata), Vimmer 1925 (LP, as punctata), Hennig 1950 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Krivosheina 1969 (L), Mamaev et al. 1977 (L), Savchenko 1986 (LP), V! (P). solatrix Osten Sacken: Alexander 1920 (LP). spec.: Crisp & Lloyd 1954 (L).Eriocera: see under Hex- atoma. Erioconopa + trivialis Meigen: Nielsen et al. 1954 (L), Okely 1979 (LP), Savchenko 1986 (L), V! (L). Erioptera (Erioptera) chlorophylla Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (LP). cladophoroides Alexander: Bruch 1939 (LP). fusculenta Edwards: Brindle 1967 (LP), Savchenko 1982 (P). flavata Westhoff: Beling 1878 (LP, as flavescens), Vimmer 1925 (LP, as flavescens), Brindle 1967 (LP, as flavescens), Savchenko 1986 (L, as geminata). + lutea Meigen: Beling 1886 (LP), Vimmer 1925 (LP), Bangerter 1930 (LP), Crisp & Lloyd 1954 (L), Lindner 1959 (L), Brindle 1967 (LP), Savchenko 1982 (L), Reusch 1988 (P). megophthalma Alexander: Alexander 1920 (LP). septemtrionis Osten Sacken: Alexander 1920 (P). squalida Loew: Houlihan 1969 (P). vespertina Osten Sacken: Alexander 1920 (P). spec.: *Dette 1916 (L, as flavescens) *Keilin 1944 (L), Peterson 1960 (L), V! (L). Erioptera (Mesocyphona) spec.: Alexander 1920 (P). Eugnophomyia + luctuosa Osten Sacken: Rogers 1928 (LP). Euphylidorea lineola Meigen: Beling 1886 (L), Levy 1919 (LP), Vimmer 1925 (LP), Brindle 1958 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (L). meigenu Verrall: Nielsen et al. 1954 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 Gy, WANG): Eutonia + barbipes Meigen: Wolff 1922 (L). marchandi Alexander: Hynes 1958 (LP). spec.: *Hennig 1950 (L, as Limnophila spec.). Geranomyia argentifera De Meijere: Alexander 1931 (P). bezzii Alexander & Leonard: Seurat 1924 (LP). caloptera Mik: Bangerter 1929 (LP, as maculipennis). canadensis Westwood: Alexander 1920 (LP), Alex- ander & Malloch 1920 (LP), Johannsen 1934 (LP). fletcheri Edwards: Alexander 1931 (LP). gelatifex Edwards: Hingston 1932 (P), Edwards 1934 (P). rostrata Say: Rogers 1927d (LP), Johannsen 1934 (L), Peterson 1960 (L), Byers 1978 (L). rubrithorax Alexander: Wood 1952 (LP). sexocellata Alexander: Wood 1952 (LP). + unicolor Haliday: Pierre 1926 (LP), Saunders 1930 (LP), Poisson 1932 (LP), Brindle 1967 (LP), Toye 1967 (L), Hinton 1968 (P). Gnophomyia jacobsoni Alexander: Rogers 1927b (LP), 1928 (L). lugubris Zetterstedt: Brindle 1967 (LP), Mamaev et al. 1977 (L, as tristis), Savchenko 1982 (LP). toschiae Alexander: Teskey 1976 (L), Alexander & Byers 1981 (L). + tristissima Osten Sacken: Malloch 1917 (LP), Alex- ander 1920 (LP), Rogers 1928 (LP). spec.: Savchenko 1986 (L). Gonempeda + flava Schummel: Brindle 1967 (P), Savchenko 1982 (P). Gonomyia (Gonomyia) kansensis Alexander: Alexander 1920 (P). lucidula De Meijere: Crisp & Lloyd, 1954 (L), Lindner 1959 (L), Brindle 1967 (LP), Savchenko, 1982 (LP). spec.: Vimmer 1925 (L), Alexander 1931 (L), Alex- ander & Byers 1981 (L), Savchenko 1986 (L). Gonomyia (Lipophleps) hawauensis Alexander: *Williams 1943 (L). pleuralis Williston: Rogers 1926 (LP). sulphurella Osten Sacken: Alexander 1920 (P), Jo- hannsen 1934 (P). sulphurelloides Alexander: Wood 1952 (LP). spec.: Alexander 1931 (L). Gonomyia (Neolipophleps) alexanderi Johnson: Alexander 1920 (LP). Gonomyodes tacoma Alexander: Hynes 1969b (LP), Byers 1978 (L). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Helius flavipes Macquart: Alexander 1920 (LP), Johannsen 1934 (P). flavus Walker: Brindle 1967 (L). + longirostris Meigen: Gercke 1884 (P), Bangerter 1929 (LP), Brindle 1967 (LP), Cramer 1968 (LP), Sav- chenko 1986 (LP), Reusch 1988 (L). mainensis Alexander: Alexander hannsen 1934 (L). pallirostris Edwards: Brindle 1967 (L). spec.: V! (L). Hesperoconopa dolichophallus Alexander: Hynes 1968 (LP). Hexatoma (Eriocera) albitarsis Osten Sacken: Alexander 1915b, 1920 (LP), Johannsen 1934 (LP). cinerea Alexander: Alexander 1920 (LP), Johannsen 1934 (LP). fultonensis Alexander: Alexander 1914a, 1920 (LP), Johannsen1934 (L), Peterson 1960 (L). longicornis Walker: Alexander 1914a, 1920 (LP). spinosa Osten Sacken: Alexander 1914a, 1920 (LP), Johannsen 1934 (L), Peterson 1960 (L), Byers 1978 (1). spec.: Malloch 1917 (L), Alexander 1931 (L), Cook 19490) NAHE Hexatoma (Hexatoma) bicolor Curtis: Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP), V! (L). fuscipennis Curtis: Brindle & Bryce 1960 (L), Brindle 1967 (LP), V! (L). | megacera Osten Sacken: Alexander 1915b (LP), 1920 (LP), Johannsen 1934 (LP), Byers 1978 (L). + nigra Latreille: *Lindner 1959 (L). spec.: *Wolff 1922 (L, as Limnophila spec.; L, as cf tenuipes), Rozkosny & Pokorny 1980 (L). Hoplolabis (Hoplolabis) + armata Osten Sacken: Alexander 1920 (P). Hoplolabis (Parilisia) vicina Tonnoir: Brindle 1967 (LP), Savchenko 1982 (LP), Reusch 1988 (P). Idioglochina: see under Dicranomyia. Idiognophomyia enneki Alexander: Byers 1974 (LP). Idioptera + pulchella Meigen: Brindle & Bryce 1960 (L), Brindle 1967 (L), Savchenko 1986 (L). Idiopyga: see under Dicranomyia. Ilisia + maculata Meigen: Crisp & Lloyd 1954 (L), Brindle 1967 (LP), Savchenko 1982 (L). Lasiomastix: see under Limnophila. Libnotes (Afrolimonia) ladogensis Lackschewitz: Savchenko 1985, 1986 (L). stantoni Edwards: * Alexander 1931 (LP). subapicalis Alexander: Wood 1952 (LP). spec.: Savchenko 1986 (L). Libnotes (Libnotes) perkinsi Grimshaw: Williams 1943 (LP). undulata Matsumura: Krivosheina 1969 (L), Mamaev et al. 1977 (L). Limnophila (Dicranophragma) + fuscovaria Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (L P). Limnophila (Elporiomyia) crepuscula Wood: Wood 1952 (LP). 1920 (LP), Jo- 257 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 + nox Alexander: Wood 1952 (LP). Limnophila (Lasiomastix) + macrocera Say: Alexander 1920 (LP), Johannsen 1934 (LP), Byers 1978 (L). Limnophila (Limnophila) (= Poecilostola) + pictipennis Meigen: Beling 1878 (LP), Brauer 1883 (LE), VI (LP). punctata Schrank: Beling 1886 (LP), Gerbig 1913 (L), Wolff 1922 (L), Brindle 1958 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Rozkosny & Pokorny 1980 (L), Savchenko 1986 (LP). spec.: Levy 1919 (LP), *Wolff 1922 (L, as spec. (= Hexatoma ?), as rufibasis), Hennig 1950 (L) (= Euto- nia?), Lindner 1959 (L), Byers 1978 (L), Savchenko 1986 (L). Limnophilomyia + Jacteitarsis Alexander: Wood 1952 (LP). Limonia flavipes Fabricius: Beling 1886 (L), Brindle 1967 (LP), Reusch 1988 (LP). hercegovinae Strobl: Reusch 1988 (L). macrostigma Schummel: Lindner 1959 (L), Brindle 1967 (LP), Savchenko 1985, 1986 (LP), Reusch 1988 (LP). nigropunctata Schummel: Beling 1878 (L). nubeculosa Meigen: Bryce 1957a (L), Lindner 1959 (L), Brindle 1967 (LP), Savchenko 1985, 1986 (LP), Reusch 1988 (LP). simulans Walker: Needham 1908 (LP), Malloch 1917 (L), Alexander 1920 (LP). + tripunctata Fabricius: Beling 1873b (LP), Lindner 1959 (L), Brindle 1967 (LP), Savchenko 1985, 1986 (LP), Reusch 1988 (LP). trivittata Schummel: V! (L). spec.: Teskey 1976 (L), Byers 1978 (L), Alexander & Byers 1981 (L). Liogma + nodicornis Osten Sacken: Alexander 1914b, 1920 (LP), Peus 1952 (LP), Brodo 1967 (LP), Alexander & Byers 1981 (L). Lipophleps: see under Gonomyia Lipsothrix errans Walker: Beling 1886 (LP, as icterica), Brindle 1967 (LP), Krivosheina 1969 (L), Savchenko 1982, 1986 (LP). fenderi Alexander: Hynes 1965 (LP). hynesiana Alexander: Hynes 1965 (LP). nigrilinea Doane: Hynes 1965 (LP). remota Walker: Hinton 1955 (P), Brindle 1967 (LP), Hinton 1967, 1968 (P). shasta Alexander: Hynes 1965 (LP). sylvia Alexander: Rogers & Byers 1956 (LP), Hynes 1965 (LP). Mesocyphona: see under Erioptera. Metalimnobia annulus cinctipes Say: Alexander 1920 (LP). bifasciata Schrank: Stannius 1829 (LP, as xanthoptera), Pastejrik 1908 (L, as xanthophora), Vimmer 1912, 1925 (L, as xanthophora), de Meijere 1917 (LP), Brauns 1954b (P), Brindle 1967 (LP), Cramer 1968 (L), Savchenko 1985 (LP), 1986 (LP), V! (LP). fallax Johnson: Alexander 1920 (LP), Johannsen 1934 (EP): immatura Osten Sacken: Malloch 1917 (P). quadrimaculata Linnaeus: Beling 1873b (LP, as annu- lus), Brauns 1954b (P), Lindner 1959 (L), Brindle 1967 258 (LP), Krivosheina 1969 (L), Savchenko 1985, 1986 (L), VI (P). quadrinotata Meigen: Lindner 1958 (LP), Brindle 1967 (LP), Savchenko 1985 (LP). triocellata Osten Sacken: Johnson 1906 (LP), Malloch 1917 (LP). Molophilus griseus Meigen: Crisp & Lloyd 1954 (L), Gachet 1967 (LP, as ripicolus), Savchenko 1982 (L). hirtipennis Osten Sacken: Alexander 1920 (LP), Byers 1978 (L). obscurus Meigen: Brindle 1967 (LP), Savchenko 1982 (P). occultus De Meijere: Okely 1979 (L), Savchenko 1986 (L). ochraceus Meigen: Beling 1886 (LP), Vimmer 1925 (LP), Brauns 1954b (P), Reusch 1988 (LP). spec.: Crisp & Lloyd 1954 (L), Lindner 1959 (L), Byers 1978 (L), Alexander & Byers 1981 (L), Savchenko 1986 (L), Winterbourn & Gregson 1989 (L). Mongoma: see under Trentepohlia. Neolimnomyia (Brachylimnophila) nemoralis Meigen: Beling 1886 (LP), Vimmer 1925 (LP), Crisp & Lloyd 1954 (L), Brindle 1958 (L), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brin- dle1967 (LP), Savchenko 1986 (LP). Neolimnomyia (Neolimnomyia) batava Edwards: Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP), Reusch 1988 (P). filata Walker: Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP). Neolimnophila carteri Tonnoir: Beling 1886 (LP, as hyalipennis). Neolimonia dumetorum Meigen: Beling 1886 (L). Neolipophleps: see under Gonomyia. Orimarga (Diotrepha) + mirabilis Osten Sacken: Rogers 1927c (LP). Orimarga (Orimarga) attenuata Walker: Vaillant 1951 (LP, as hygropetrica), Savchenko 1985 (LP). Orimargula: see under Antocha.Ormosia hederae Curtis: V! (P). lineata Meigen: Lindner 1959 (L), Brindle 1967 (L), Savchenko 1982, 1986 (L). meigenti Osten Sacken: Alexander 1920 (LP), Byers 1978 (L). nigripila Osten Sacken: Alexander 1920 (P). romanovichiana Alexander: Alexander 1920 (LP, as nubila). spec.: Johannsen 1934 (L), Byers 1978 (L), Alexander & Byers 1981 (L), Savchenko 1982 (LP), Reusch 1988 (P). Oxyrhiza: see under Paradelphomyia. Paradelphomyia (Oxyrhiza) (= Oxydiscus). americana Alexander: * Alexander 1920 (L). cayuga Alexander: *Alexander 1920 (L). minuta Alexander: *Alexander 1920 (LP), Johannsen 1934 (L), Savchenko 1986 (P). senilis Haliday: Crisp & Lloyd 1954 (L), Lindner 1959 (L), Brindle 1967 (L), Krivosheina 1969 (L), Sav- chenko 1986 (L), Reusch 1988 (P). spec.: Brindle & Bryce 1960 (L). Paradicranota: see under Dicranota. Paralimnophila skusei Hutton: Winterbourn & Gregson 1989 (L). Paramongoma: see under Trentepohlia. Paraphylidorera: see under Phylidorea. Parilisia: see under Ilisia. Pedicia (Amalopis) + occulta Meigen: Lindner 1959 (L), Brindle 1967 (L), Savchenko 1986 (L). Pedicia (Crunobia) littoralis Meigen: Brindle 1967 (L). + straminea Meigen: Beling 1878 (LP, as schineri), Vimmer 1925 (LP, as schineri), Lindner 1959 (L), Brindle 1967 (L), Savchenko 1986 (L). spec.: Brindle 1962 (L). Pedicia (Pedicia) albivitta Walker: Needham 1903 (L, as unidentified), 1908 (L), Alexander 1920 (L), Johannsen 1934 (L), Byers 1978 (L). + rivosa Linnaeus: Beling 1878 (LP), Levy 1919 (LP), Vimmer 1925 (LP), Oldham 1926 (L), Wardle 1926 (L), Wardle & Taylor 1926 (L), Wesenberg-Lund 1943 (L), Hennig 1950 (L), Brauns 1954b (P), Crisp & Lloyd 1954 (L), Brindle 1962 (L), 1967 (LP), Sav- chenko 1986 (LP), V! (LP). spec.: Alexander & Byers 1981 (L). Phalacrocera + replicata Linnaeus: Bengtsson 1897 (L), Miall & Shel- ford 1897 (LP), Malloch 1917 (LP), Alexander 1920 (LP), Lenz 1920b (LP), Haake 1922 (LP), Johannsen 1934 (LP), Wesenberg-Lund 1943 (L), Peus 1952 (LP), Brindle 1967 (LP), Brodo 1967 (LP), Krivo- sheina 1969 (L), Byers 1978 (L), V! (L). tipulina Osten Sacken: Brodo 1967 (P). vancouverensts Alexander: Spencer 1930 (LP). Phylidorea (Macrolabina) + nigronotata Siebke: Wiedenska 1987 (LP). Phylidorea (Paraphylidorea) + fulvonervosa Schummel: Crisp & Lloyd 1954 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Sav- chenko 1986 (L), Wiedenska 1987 (LP), Reusch 1988 (P). Phylidorea (Phylidorea) abdominalis Staeger: Brindle 1967 (LP). adusta Osten Sacken: *Alexander 1920 (LP), Jo- hannsen 1934 (P). + ferruginea Meigen: Beling 1878 (P, as lineola), de Meijere 1917 (LP), *Levy 1919 (LP), Wolff 1922 (L), Hennig 1950 (P), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Savchenko 1986 (LP), V! (P). longicornis Schummel: Brindle 1967 (P, as glabricula). nervosa Schummel: Wiedenska 1987 (LP). squalens Zetterstedt: Brindle & Bryce 1960 (L), Brin- dle 1967 (LP), Savchenko 1986 (LP), Wiedenska 1987 (LP). Pilaria discicollis Meigen: Gerbig 1913 (L), Levy 1919 (LP), Vimmer 1924, 1925 (P), Brindle 1958 (L), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brindle 1967 (LP), Rozkosny & Pokorny 1980 (L), Savchenko 1986 (LP), Reusch 1988 (LP), V! (L). fuscipennis Meigen: Brauer 1883 (L), Beling 1886 (LP), Vimmer 1906 (L), Gerbig 1913 (L), Levy 1919 (LP), Wolff 1922 (L), Vimmer 1925 (LP), Rozkosny & Pokorny 1980 (L), V!(L). quadrata Osten Sacken: Alexander 1920 (P). recondita Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (L), Alexander & Byers 1981 (L). OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea ripicola Gérard: Gérard 1968 (LP). scutellata Staeger: Brindle & Bryce 1960 (L), Brindle 1967 (LP). tenuipes Say: Malloch 1917 (L), Alexander 1920 (LP), Johannsen 1934 (LP). spec.: Anton 1943a (L, as ? Ulomorpha), Hennig 1950 (L, as ? Ulomorpha), Crisp & Lloyd 1954 (L), Byers 1978 (L). Polymera georgiae Alexander: Rogers 1933 (LP). rogerstana Alexander: Rogers 1933 (LP). Progonomyia: see under Ellipteroides. Protogonomyia: see under Ellipteroides. Pseudolimnophila inornata Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (P), Byers 1978 (L), Alexander & Byers 1981 (L). lucorum Meigen: Crisp & Lloyd 1954 (L), Hinton 1954 (P), Brindle 1958 (L), Brindle & Bryce 1960 (L), Brin- dle 1967 (LP), Rozkosny & Pokorny 1980 (L), Sav- chenko 1986 (LP), Reusch 1988 (P). + luteipennis Osten Sacken: Hart 1895 (LP), Malloch 1917 (LP), Alexander 1920 (LP), Johannsen 1934 (L), V! (L). Psiloconopa: see under Symplecta. Rhabdomastix (Sacandaga) afra Wood: Wood 1952 (LP). californiensis Alexander: Hynes 1969c (LP). fascigera Alexander: Hynes 1969c (LP). flaviventris Alexander: Hynes 1969c (LP). , hynesi Alexander: Hynes 1969c (LP). leonardi Alexander: Hynes 1969c (LP). schistacea Schummel: Beling 1886 (LP), Vimmer 1925 (LP). setigera Alexander: Hynes 1969c (LP), Byers 1978 (L). subfascigera Alexander: Hynes 1969c (LP). trichophora Alexander: Hynes 1969c (LP). spec.: Savchenko 1982 (LP). Rhamphophila + obscuripennis Hudson: Hudson 1920 (LP, sinistra). Rhaphidolabina: see under Dicranota. Rhaphidolabis: see under Dicranota. Rhipidia bryanti Johnson: Alexander 1920 (LP), Johannsen 1934 (LP). + maculata Meigen (= duplicata authors): Beling 1873b (P), 1878 (L), Brauns 1954b (P), Brindle 1967 (L), Savchenko 1985 (L), Reusch 1988 (LP). uniseriata Schiner: Beling 1878 (L). Rhypholophus haemorrhoidalis Zetterstedt: Beling 1878 (P), 1886 (LP), Crisp & Lloyd 1954 (L), Lindner 1959 (L), Brin- dle 1967 (LP), Krivosheina 1969 (L), Savchenko 1982, 1986 (LP), V! (P). varius Meigen: de Meijere 1917 (LP), Wolff 1922 (L), Brindle 1967 (L P). spec. V! (LP). Sacandaga: see under Rhabdomastix. Scleroprocta innocens Osten Sacken: Alexander, 1920 (P), Jo- hannsen 1934 (P), Brindle 1967 (P). + sororcula Zetterstedt: Brindle 1967 (LP, as danica). spec.: Alexander 1920: Figs 405-406 (as unidentified), Bangerter 1930 (LP, as fascipennis), Savchenko 1982 (LP). 259 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Sigmatomera (Austrolimnobia) rufa Hudson: Hudson 1920 (LP). Sigmatomera (Sigmatomera) shannoniana Alexander: Alexander 1930 (L). Styringomyia ingrami Edwards: Edwards 1924 (P). neocaledoniae Alexander: Hynes 1990 (LP). Symplecta (Psiloconopa) graphica Osten Sacken: Hart 1895 (L). stictica Meigen: Brindle 1967 (LP), Houlihan 1969 (P), Reusch 1988 (LP). Symplecta (Symplecta) cana Walker: Hart 1895 (LP, as punctipennis), *Mal- loch 1917 (LP, as punctipennis), Vimmer 1925 (LP, as punctipennis). + hybrida Meigen: Beling 1878 (LP, as punctipennis), Alexander 1920 (LP), Johannsen 1934 (LP), Sav- chenko, 1982 (LP), Reusch 1988 (LP), V! (L). macroptera Philippi: Bruch 1939 (LP). Symplecta (Trimicra) + pilipes Fabricius: Beling 1878 (LP), Gerbig 1913 (L), Alexander 1920 (L), Pierre 1924 (P, as marina), Vimmer 1925 (LP), Johannsen 1934 (L), Bruch 1939 (LP), Séguy 1942 (LP, as pazliana), Wood 1952 (LP, as inconspicua), Vaillant 1953 (LP, as hirsutipes), Brindle 1967 (L), Byers 1978 (L), Savchenko 1982 (L). Teucholabis + complexa Osten Sacken: Alexander 1920 (LP). Thaumastoptera + calceata Mik: Lenz 1920a (LP), Liang 1925 (LP), Brauns 1954b (P), Vaillant 1956 (L), Brindle 1967 (LP), Savchenko 1985 (LP). ; spec.: Vimmer 1929 (L). Tonnoiraptera + neozelandica Tonnoir: Tonnoir 1926 (P). Trentepohlia (Mongoma) pennipes Osten Sacken: de Meijere 1911 (LP), Alex- ander 1920 (LP). Trentepohlia (Paramongoma) bromeliadicola Alexander: Picado 1913 (LP), Alex- ander 1920 (LP). Tricyphona + immaculata Meigen: Beling 1878 (L), de Meijere 1917 (LP), Lindner 1959 (L), Brindle 1962 (L), 1967 (LP), Okely 1979 (L), Savchenko 1986 (LP). inconstans Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (L), Byers 1978 (L). schummeli Edwards: Brindle 1967 (L). unicolor Schummel: Brindle 1962 (L), 1967 (LP). Trimicra: see under Symplecta. Triogma exsculpta Osten Sacken: Brodo 1967 (LP). + trisulcata Schummel: Alexander 1920 (LP), Lenz 1920b (LP), Haake 1922 (LP), Johannsen 1934 (L), Wesenberg-Lund 1943 (L), Peus 1952 (LP), Brindle 1967 (LP), V! (L). Ula bolitophila Loew: Krivosheina 1969 (L), Krivosheina et al. 1986 (L), Savchenko 1986 (L). elegans Osten Sacken: Alexander 1915a (LP), Malloch 1917 (LP), Alexander 1920 (LP). + mollissima Haliday: Engel 1916 (L, as macroptera), Vimmer 1925 (LP, as macroptera), Lindner 1959 (L, as crassicauda), Savchenko 1986 (L). sylvatica Meigen: Bangerter 1934 (LP), Bryce 1957b (L), Lindner 1959 (L), Brindle & Bryce 1960 (L), Brin- 260 dle 1967 (LP), Savchenko 1986 (LP), V! (LP). — Immature stages of Japanese Ula species are described by Tokunaga et al. 1954 (not consulted). Ulomorpha + pilosella Osten Sacken: Alexander 1920 (LP), Jo- hannsen 1934 (L). spec. Anthon 1943a (L), Hennig 1950 (L). 11. ACKNOWLEDGEMENTS The authors would like to thank G. W. Byers (Lawrence) and J. K. Gelhaus (Philadelphia) for making available larval material of several genera of Tipulidae, B. Sinclair (Ottawa) for providing larvae of Dactylolabis and valuable notes on the distribution of the pharyngeal filter apparatus, H. de Jong (Amsterdam) for his valuable comments on a draft of this manuscript, and W. Langerak (Amsterdam) who has redrawn most figures. 12. REFERENCES Alexander, C. P., 1914a. The biology of the North Amer- ican crane flies (Tipulidae, Diptera). 1. The genus Eriocera Macquart. - Pomona College Journal of En- tomology and Zoology 6: 12-34. Alexander, C. P., 1914b. Biology of the North American crane flies (Tipulidae, Diptera). 2. Liogma nodicornis Osten Sacken. - Pomona College Journal of Entomol- ogy and Zoology 6: 105-118. Alexander, C. P., 1915a. The biology of the North Amer- ican crane flies (Tipulidae, Diptera). 3. The genus Ula Haliday. - Pomona College Journal of Entomology and Zoology 7: 1-8. Alexander, C. P., 1915b. The biology of the North Amer- ican crane flies (Tipulidae, Diptera). 4. The tribe Hex- atomini. - Pomona College Journal of Entomology and Zoology 7: 141-158. Alexander, C. P., 1919a. The crane flies of New York, Part 1. Distribution and taxonomy of the adult flies. - Cornell University Agricultural Experimental Station Memoirs 25: 771-993. Alexander, C. P., 1919b. The biology of the North Amer- ican crane flies (Tipulidae, Diptera). 5. The genus Dicranoptycha Osten Sacken. - Pomona College Jour- nal of Entomology and Zoology 11: 67-74. Alexander, C. P., 1920. The crane flies of New York, Part 2. Biology and phylogeny. - Cornell University Agri- cultural Experimental Station Memoirs 38: 691-1133. Alexander, C. P., 1922. The biology of the North Amer- ican crane flies (Tipulidae, Diptera). 6. The genus Cladura Osten Sacken. - Pomona College Journal of Entomology and Zoology 14: 1-6. Alexander, C. P., 1927. The interpretation of the radial field of the wing in the nematocerous Diptera, with special reference to the Tipulidae. - Proceedings of the Linnean Society of New South Wales 52: 41-72. Alexander, C. P., 1930. The genus Sigmatomera Osten Sacken with observations on the biology by Raymond C. Shannon. - Encyclopédie Entomologique, B II, Dip- tera 5: 155-162. Alexander, C. P., 1931. Deutsche Limnologische Sunda- Expedition. The crane flies (Tipulidae, Diptera). - Archiv fiir Hydrobiologie, Suppl. Band 9: Tropische Binnengewässer 2: 135-191. Alexander, C. P., 1972. Diptera: Tipulidae. - Insects of Micronesia 12: 733-863. Alexander, C. P. & G. W. Byers, 1981. Tipulidae. - In: Manual of Nearctic Diptera, volume 1. (edited by J. F. McAlpine et al.): 153-190, Biosystematic Research Institute, Ottawa, Ontario, Monograph 27: i-vi, 1-674. Alexander, C. P. & J. R. Malloch, 1920. Notes on the life history of a crane fly of the genus Geranomyia Haliday (Tipulidae, Diptera). - Transactions of the Illinois State Academy of Science 13: 310-319. Anthon, H., 1943a. Der Kopfbau der Larven einiger nematoceren Dipterenfamilien. — Spolia Zoologica Musei Hauniensis 3: 1-61, 11 pl. Anthon, H., 1943b. Zum Kopfbau der primitivsten bisher bekannten Dipterenlarve: Olbiogaster sp. (Rhyphidae). Ein Beitrag zur Phylogenie der nemat- oceren Dipteren. - Entomologiske Meddelelser 23: 303-320. Anthon, H., 1988. Larval morphology of Mischoderus (Insecta, Diptera, Nematocera, Tanyderidae) with notes on tanyderid affinities. - Zoologica Scripta 17: 381-397. Bangerter, H., 1928. Miicken-Metamorphosen. 1. - Ko- nowia 7: 156-161. Bangerter, H., 1929. Mücken-Metamorphosen. 2. - Ko- nowia 8: 1-6. Bangerter, H., 1930. Mücken-Metamorphosen, 3. - Ko- nowia 9: 97-102. Bangerter, H., 1931. Mücken-Metamorphosen. 4. - Ko- nowia 10: 191-196. Bangerter, H., 1934. Miicken-Metamorphosen. 6. - Ko- nowia, 13: 264-272. Beling, T., 1873a. Acht neue Arten Deutscher zweiflüge- liger Insecten. - Verhandlungen der Zoologischen- Botanischen Gesellschaft in Wien 23: 547-560. Beling, T., 1873b. Beitrag zur Naturgeschichte verschied- ener Arten aus der Familie der Tipuliden. - Verhand- lungen der Zoologischen-Botanischen Gesellschaft in Wien 23: 575-592. Beling, T., 1878. Zweiter Beitrag zur Naturgeschichte (Metamorphose) verschiedener Arten aus der Familie der Tipuliden. - Verhandlungen der Zoologischen- Botanischen Gesellschaft in Wien 28: 21-56. Beling, T., 1886. Dritter Beitrag zur Naturgeschichte (Metamorphose) verschiedener Arten aus der Familie der Tipuliden. - Verhandlungen der Zoologischen- Botanischen Gesellschaft in Wien 36: 171-214. Bengtsson, S., 1897. Bidrag till kinnedomen om larven af Phalacrocera replicata (Lin.). — Acta Universitatis Lundensis 33 (7): 1-102. Brauer, F., 1883. Die Zweiflügler des kaiserlichen Muse- ums in Wien. 3. - Kaiserlich-Königlichen Hof- und Staatsdruckerei, Wien: 100 pp, 5 pl. Brauns, A., 1954a. Terricole Dipterenlarven. - Mus- terschmidt, Göttingen: 179 pp, 96 figs. Brauns, A., 1954b. Puppen terricoler Dipterenlarven. - Musterschmidt, Göttingen: 156 pp, 75 figs. Brindle, A., 1958. Notes on the identification of Limno- phila larvae (Diptera-Tipulidae). 1. - Transactions of the Society for British Entomology 13: 57-68. Brindle, A., 1960. The larvae and pupae of the British Tipulinae. - Transactions of the Society for British Entomology 14: 63-114. Brindle, A., 1962. The natural groups of the British Pe- diciini (Dipt., Tipulidae). - Entomologist’s Monthly OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Magazine 98: 234-237. Brindle, A., 1967. The larvae and pupae of the British Cylindrotominae and Limoniinae (Diptera, Tipuli- dae). - Transactions of the Society for British Ento- mology 17: 151-216. Brindle, A. & D. Bryce, 1960. The larvae of the British Hexatomini (Dipt., Tipulidae). — Entomologist’s Gazette 11: 207-224. Brodo, F, 1967. A review of the subfamily Cylindrotomi- nae in North America (Diptera: Tipulidae). - Univer- sity of Kansas Science Bulletin 48: 71-115. Bruch, C., 1939. Contributién al conocimiento de los ti- pulidos Argentinos (Diptera). - Physis, 17: 3-28. Bryce, D., 1957a. Notes on the life history of British Limoniinae (Dipt., Tipulidae). 1. Limonia nubeculosa Meig. and L. didyma Meig. - Entomologist’s Monthly Magazine 93: 67-69. Bryce, D., 1957b. Notes on the life history of British Limoniinae (Dipt., Tipulidae). 2. Ula sylvatica Mei- gen. - Entomologist's Monthly Magazine 93: 132. Byers, G. W., 1961. The crane fly genus Dolichopeza in North America. - Kansas University Science Bulletin 42: 665-924. Byers, G. W., 1963. The life history of Panorpa nuptialis (Mecoptera: Panorpidae). - Annals of the Entomo- logical Society of America 56: 142-149. Byers, G. W., 1974. Larva and pupa of Idiognophomyia enniki Alexander (Diptera: Tipulidae). - Pan-Pacific Entomologist 50: 282-287. Byers, G. W., 1978. Tipulidae. - In: An introduction to the aquatic insects of North America (edited by R. W. Merritt & K. W. Cummins): 285-310, Kendall & Hunt, Dubuque (Ohio). Byers, G. W., 1983. The crane fly genus Chionea in North America. - Kansas University Science Bulletin 52: 59- 195. Cameron, A. E., 1918. Life history of the leaf eating crane- fly Cylindrotoma splendens Doane. - Annals of the Entomological Society of America 9: 67-89. Chiswell, J. R., 1955. On the last instar larva of Tipula livida van der Wulp (Diptera, Tipulidae) with notes on the fronto-clypeal region of larval Tipulinae and caterpillars. - Proceedings of the Royal Entomological Society of London (A) 30: 127-136. Cook, E. F., 1949. The evolution of the head in the larvae of the Diptera. - Microentomology 14: 1-57. Courtney, G. W., 1990. Cuticular morphology of larval mountain midges (Diptera: Deuterophlebiidae): im- plications for the phylogenetic relationships of Nematocera. - Canadian Journal of Zoology 68: 556- 578. Courtney, G. W., 1991. Phylogenetic analysis of the Ble- phariceromorpha, with special reference to mountain midges (Diptera: Deuterophlebiidae). — Systematic Entomology 16: 137-172. Cramer, E., 1968. Die Tipuliden des Naturschutzparkes Hoher Vogelsberg. — Deutsche Entomologische Zeitschrift 15: 133-232. Crisp, G. & L. Lloyd, 1954. The community of insects in a patch of woodland mud. - Transactions of the Royal Entomological Society of London 105: 269-313. Dahl, C., 1980. Comparison of postembryonic organiza- tion of the genital segments in Trichoceridae, Tipuli- dae, and Anisopodidae (Diptera, Nematocera). — Zoo- logica Scripta 9: 165-185. Dahl, C. & C. P. Alexander, 1976. A world catalogue of 261 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Trichoceridae Kertész, 1902 (Diptera). - Entomolog- ica Scandinavica 7: 7-18. Dette, E., 1916. Uber die Metamorphose von Trichosti- cha flavescens. — Zoologische Jahrbücher, Abteilung fiir Systematik, Okologie und Geographie der Tiere 39: 417-439. Dienske, J. W., 1987. An illustrated key to the genera and subgenera of the western palaearctic Limoniidae (In- secta: Diptera), including a description of the external morphology. - Stuttgarter Beiträge zur Naturkunde (A) 409: 1-52. Edwards, F. W., 1924. New data concerning Styringomyia (Diptera, Tipulidae). - Annals and Magazine of Nat- ural History (9) 13: 265-274. Edwards, F. W., 1926. The phylogeny of nematocerous Diptera: a critical review of some recent suggestions. — Verhandlungen des III. Internationalen Entomolo- gen-Kongresses 1: 111-130. Edwards, F. W., 1928. Diptera. Fam. Protorhyphidae, Anisopodidae, Pachyneuridae, Trichoceridae, with descriptions of early stages by D. Keilin. - Genera Insectorum 190: 41 pp, 2 pl. Edwards, F. W., 1934. Oxford University expedition to British Guiana, 1929. Diptera Nematocera. - Annals and Magazine of Natural History (10) 14: 632-635. Edwards, F. W., 1936. Pupae and living adults of Chionea lutescens Lundström from Denmark. - Proceedings of the Royal Entomological Society of London 10: 89-91. Egger, J., G. Frauenfeld & F. Brauer, 1854. Beobachtungen über die Entwicklungsgeschichte der Chionea ara- neoides, nebst Anatomie des Insectes und der Larve. - Verhandlungen des Zoologischen-Botanischen Ve- reins in Wien 4: 609-616. Engel, E. O., 1916. Beitrige zur Kenntnis einiger Dipter- enlarven. - Mitteilungen der Münchner Entomolog- ischen Gesellschaft 7: 68-76. Fahy, E, 1972. The larva and life history of Dicranota guerini Zett. (Dipt. Tipulidae). - Entomologist 105: 260-263. Gachet, M., 1967. Un diptere Limoniidae nouveau et ses premiers stades. - Travaux du Laboratoire d’Hydrobi- ologie et Pisciculture de Grenoble 57-58: 51-55. Gelhaus, J. K., 1986. Larvae of the crane fly genus Tipula in North America (Diptera: Tipulidae). - Kansas University Science Bulletin 53: 121-182. Gelhaus, J. K. & C. W. Young, 1991. The immature stages and biology of the crane fly genus Brachypremna (Diptera: Tipulidae). - Proceedings of the Entomo- logical Society of Washington (in press). Gérard, M., 1968. Pilaria ripicola n. sp. (Diptera Limonii- dae). Morphologie de la larve, de la nymphe et de imago. Anatomie de la larve. - Travaux du Labora- toire d’ Hydrobiologie et Pisciculture de Grenoble 59- 60: 165-184. Gerbig, F, 1913. Uber Tipulidenlarven mit besonderer Berücksichtigung der Respirationsorgane. - Zoolo- gische Jahrbücher, Abteilung für Systematik, Geogra- phie und Biologie der Tiere 35: 127-181. Gercke, G., 1884. Über de Metamorphose einiger Dip- teren. - Verhandlungen des Vereins für Naturwissen- schaftliche Unterhaltung zu Hamburg 5: 68-80. Giljarov, M. C. (Ed.), 1964. Keys to the grounddwelling insect larvae. - Nauk, Moscow: 919 pp. (in Russian). Goetghebuer, M., 1925. Contribution à l'étude des ‘pré- mandibules’ chez les larves des Diptères nématocères. 262 — Encyclopédie Entomologique, B II, Diptera 1: 142- 1522 Griffiths, G. C. D., 1990. Review of: McAlpine, J. E. et al. (Eds). Manual of Nearctic Diptera, Vol. 3. - Quaesti- ones Entomologicae 26: 117-130. Haake, B., 1922. Die Metamorphose von Triogma trisul- cata. - Zoologische Jahrbücher, Abteilung für Syste- matik, Geographie und Biologie der Tiere 45: 459- 500. Harbach, R. E. & K. L. Knight, 1980. Taxonomist’s glos- sary of mosquito anatomy. - Department of Entomol- ogy, North Carolina State University, Raleigh: 12 + 415 pp. Hart, C. A., 1895. On the entomology of the Illinois river and adjacent waters. First Paper. - Bulletin of the Illinois State Laboratory of Natural History 4: 149- Di Hemmingsen, A. M., B. Mannheims & P. Nielsen, 1959. Notes on crane-flies (Tipulidae) collected by Dr. Axel M. Hemmingsen in the Canary Islands. - Entomolo- giske Meddelelser 29: 71-77. Hennig, W., 1948. Die Larvenformen der Dipteren, Vol. 1. - Akademie-Verlag, Berlin: 185 pp. Hennig, W., 1950. Die Larvenformen der Dipteren, Vol. 2.- Akademie-Verlag, Berlin: 8 + 458 pp. Hennig, W., 1952. Die Larvenformen der Dipteren, Vol. 3. - Akademie-Verlag, Berlin: 8 + 628 pp. Hennig, W., 1954. Flügelgeäder und System der Dipteren. - Beiträge zur Entomologie 4: 245-388. Hennig, W., 1966. Phylogenetic systematics (Translated by D. D. Davis & R. Zangerl). - University of Illinois Press, Urbana: 16 + 263 pp. Hennig, W., 1968. Kritische Bemerkungen über den Bau der Flügelwurzel bei den Dipteren und die Frage nach der Monophylie der Nematocera. — Stuttgarter Bei- träge zur Naturkunde 195: 1-23. Hennig, W., 1973. Diptera. - Handbuch der Zoologie, 4 (2)72/31:1-3372 Hennig, W., 1981. Insect Phylogeny (Translated and edited by A. C. Pont with revisionary notes organized by D. Schlee). - J. Wiley & Sons, New York: 22 + 514 PP- Hingston, R. W. G., 1932. A naturalist in the Guiana forest. - Arnold, London:16 + 384 pp. Hinton, H. E., 1947. On the reduction of functional spir- acles in the aquatic larvae of the Holometabola, with notes on the moulting process of spiracles. - Transac- tions of the Royal Entomological Society of London 98: 449-473. Hinton, H. E., 1950. Aquatic Diptera collected in the river Dove near Dovedale, Derbyshire. - Journal of the Society for British Entomology 3: 203-206. Hinton, H. E., 1954. On the structure and function of the respiratory horns of the pupae of the genus Pseudo- limnophila (Diptera: Tipulidae). - Proceedings of the Royal Entomological Society of London, (A) 39: 135- 140. Hinton, H. E., 1955. The structure of the spiracular gill of the genus Lipsothrix (Tipulidae), with some obser- vations on the living epithelium isolated in the gill at the pupa-adult moult. - Proceedings of the Royal Entomological Society of London, (A) 30: 1-14. Hinton, H. E., 1957. The structure and function of the spiracular gill of the fly Taphrophila vitripennis. - Proceedings of the Royal Society of London, (B) 147: 90-120. Hinton, H. E., 1965. The spiracular gill of the fly Orimar- gula australiensis and its relation to those of other insects. — Australian Journal of Zoology 13: 783-800. Hinton, H. E., 1966. The spiracular gill of the fly Antocha bifida, as seen with the scanning electron microscope. — Proceedings of the Royal entomological Society of London, (A) 41: 107-115. Hinton, H. E., 1967. Structure of the plastron in Lipso- thrix, and the polyphyletic origin of plastron respira- tion in Tipulidae. - Proceedings of the Royal Entomo- logical Society of London, (A) 42: 35-38. Hinton, H. E., 1968. Spiracular gills. - Advances in Insect Physiology 5: 65-161. Houlihan, D. FE, 1969. The structure and behaviour of Notiphila riparia and Erioptera squalida, two root- piercing insects. - Journal of Zoology, London 159: 249-267. Hudson, G. V., 1920. Illustrated life-histories of New Zealand insects.1. - Transactions and Proceedings of the New Zealand Institute 52: 32-34. Hutson, A. M., 1980. Family Tipulidae. - In: Catalogue of the Diptera of the Afrotropical Region (edited by R. W. Crosskey): 47-91, British Museum (Natural His- tory), London. Hynes, C. D., 1958. A description of the immature stages of Limnophila (Eutonia) marchandi Alex. (Diptera, Tipulidae). - Proceedings of the Entomological So- ciety of Washington 60: 9-14. Hynes, C. D., 1963. Description of the immature stages of Cryptolabis magnistyla Alexander (Diptera, Tipu- lidae). - Pan-Pacific Entomologist 39: 255-260. Hynes, C. D., 1965. The immature stages of the genus Lipsothrix in the western United States (Diptera, Tipulidae). - Pan-Pacific Entomologist 41: 165-172. Hynes, C. D., 1968. The immature stages of Hesperocon- opa dolichophallus (Alex.) (Diptera: Tipulidae). - Pan-Pacific Entomologist 44: 324-327. Hynes, C. D., 1969a. The immature stages of Arctocon- opa carbonipes (Alex.) (Diptera: Tipulidae). - Pan- Pacific Entomologist 45: 1-3. Hynes, C. D., 1969b. The immature stages of Gono- myodes tacoma Alex. (Diptera: Tipulidae). - Pan- Pacific Entomologist 45: 116-119. Hynes, C. D., 1969c. The immature stages of the genus Rhabdomastix (Diptera: Tipulidae). - Pan-Pacific En- tomologist 45: 229-237. Hynes, C. D., 1990. Description of the immatures of Styringomyia neocaledoniae Alexander (Diptera: Ti- pulidae) and notes on its biology. - Pan-Pacific Ento- mologist 66: 89-92. Johannsen, O. A., 1934. Aquatic Diptera. 1. Nemocera, exclusive of Chironomidae and Ceratopogonidae. - Cornell University Agricultural Experimental Station Memoirs 164: 1-70. Johnson, C. W., 1906. Notes on some dipterous larvae. — Psyche 13: 1-4. Kaltenbach, A., 1978. Mecoptera. - Handbuch der Zool- ogie, 4 (2) 2/28: 1-111. Kawada, A., 1960. Illustrated insect larvae of Japan (3 ed.). - Hokuryukan, Tokyo:712 + 73 + 50 pp, 4 pl. Keilin, D. ,1912. Recherches sur les Diptères du genre Trichocera. Biologie, Développement, Affinités. — Bulletin Scientifique de la France et de la Belgique (7) 46: 172-191 Keilin, D., 1919. On the structure of the larvae and the systematic position of the genera Mycetobia Mg., Di- OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea tomyia Winn., and Symmerus Walk. (Diptera, Nemat- ocera). — Annals and Magazine of Natural History (9) 3: 33-42. Keilin, D., 1944. Respiratory systems and respiratory adaptations in larvae and pupae of Diptera. — Parasi- tology 36: 1-66. Keilin, D. & P. Tate, 1940. The early stages of the families Trichoceridae and Anisopodidae (+Rhyphidae) (Dip- tera: Nematocera). — Transactions of the Royal Ento- mological Society of London 90: 39-62. Krivosheina, N. P., 1964. Families Tipulidae, Limoniidae, Cylindrotomidae. — In: Keys to the grounddwelling insect larvae (edited by M. C. Giljarov): 665-710. Nauk, Moscow (in Russian). Krivosheina, N. P., 1969. Ontogenesis and evolution of the Diptera. - Nauk, Moscow, 289 pp (in Russian). Krivosheina, N. P. & B. M. Mamaev, 1970. The family Cramptonomyiidae (Diptera, Nematocera), new for the fauna of the U. S. S. R., its morphology, ecology and phylogenetic relationships. - Entomologicheskoe Obozrenie 49: 886-898 (in Russian; English version in Entomological Review, 49: 541-548). Krivosheina, N. P., A. I. Zaitzev & E. B. Jakovlev, 1986. Insects damaging forest fungi in the European part of the USSR. - Nauk, Moscow: 309 pp (in Russian). Lackschewitz, P., 1932. Eine neue Elephantomyia O. S. (Dipt., Nematoc. polyn.) aus dem Ostbaltikum. - Ko- nowia 11: 218-222 Lackschewitz, P. & F. Pagast, 1940. 16. Limoniidae. - Fliegen der Palaearktischen Region, 3(5)2, Lief. 135: 1-16. Lackschewitz, P. & F. Pagast, 1941. 16. Limoniidae. - Fliegen der Palaearktischen Region, 3(5)2, Lief. 139: 17-32. Lackschewitz, P. & F. Pagast, 1942. 16. Limoniidae. - Fliegen der Palaearktischen Region, 3(5)2, Lief. 145: 33-64. Laurence, B. R., 1953. The larva of Ectaetia (Dipt., Sca- topsidae). — Entomologist's Monthly Magazine 89: 204-205. Lenz, E, 1920a. Thaumastoptera calceata Mik. Eine ge- häusetragende Tipulidenlarve. - Archiv für Naturge- schichte 85 (A) 4: 114-136. Lenz, F., 1920b. Die Metamorphose der Cylindrotomiden. - Archiv für Naturgeschichte 85 A (6): 113-146. Levy, L., 1919. Contributions a l'étude des métamor- phoses aquatiques des diptères. - Annales de Biologie Lacustre 9: 201-248. Liang, S., 1925. Morphologie des Hypopygiums, der män- nlichen Genitaldriisen und des Verdauungssystems von Thaumastoptera calceata Mik (Tipulidae, Dip- tera). — Archiv fiir Naturgeschichte, Abt. A 41: 1-31. Lindner, Eb., 1958. Pilz bewohnende Limoniiden larven unter besonderer beruecksichtigung von Limonia quadrinotata Meigen (Diptera). - Tijdschrift voor En- tomologie 101: 263-281. Lindner, Eb., 1959. Beiträge zur Kenntnis der Larven der Limoniidae. - Zeitschrift für Morphologie und Ökol- ogie der Tiere 48: 209-319. Lindner, Er., 1949. Handbuch. - Fliegen der palaearkti- schen Region 1: 1-422. Maddison, W. P., M. J. Donoghue & D. R. Maddison, 1984. Outgroup analysis and parsimony. — Systematic Zool- ogy 33: 83-103. Malloch, J. R., 1917. A preliminary classification of Dip- tera, exclusive of Pupipara, based upon larval and 263 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 pupal characters, with keys to imagines in certain families. 1. - IIlinois State Laboratory of Natural His- tory Bulletin 12: 161-409. Mamaev, B. M. & N. P. Krivosheina, 1965. Diptera, Ce- cidomyiidae. - Nauk, Moscow: 278 pp (in Russian). Mamaev, B. M. & N. P. Krivosheina, 1966. New data on the taxonomy and biology of Diptera of the family Axymyiidae. — Entomologicheskoe Obozrenie 45: 168-180 (in Russian; English version in Entomolog- ical Review, 45: 93-99). Mamaev, B. M., N. P. Krivosheina & V. A. Potockaja, 1977. Classification key of the larvae of predatory insects-entomophages of trunk pests. - Nauk, Mos- cow: 392 pp (in Russian). Matsuda, R., 1965. Morphology and evolution of the insect head. - Memoirs of the American Entomolog- ical Institute 4: 1-334. McAlpine, J. F. (Ed.), 1981. Manual of Nearctic Diptera, Vol. 1. - Biosystematic Research Institute, Ottawa, Ontario, Monograph 27: i-vi, 1-674. McAlpine, J. F. (Ed.), 1989. Manual of Nearctic Diptera, Vol. 3. - Biosystematic Research Centre, Ottawa, On- tario, Monograph 32: i-vi, 1333-1581. De Meijere, J. C. H., 1911. Studien über südostasiatischen Dipteren. V. Ostindische Tipuliden. - Tijdschrift voor Entomologie 54: 21-79. De Meijere, J. C. H., 1917. Beiträge zur Kenntnis der Dipteren-Larven und -Puppen. - Zoologische Jah- rbücher, Abteilung für Systematik, Okologie und Geographie der Tiere 40: 177-332. Miall, L. C., 1893. Dicranota; a carnivorous Tipulid larva. - Transactions of the Entomological Society of Londen 1893: 235-253. Miall, L. C., 1934: The natural history of aquatic insects. — MacMillan, London: 11 + 395 pp. Miall, L. C. & R. Shelford, 1897. The structure and life history of Phalacrocera replicata. - Transactions of the Entomological Society of Londen 1897: 343-361. Mik, J., 1884. Biologische Fragmente. 1. Die Nymphe von Trochobola caesarea O.S. (Diptera). - Wiener Ento- mologische Zeitung 3: 65-71. Mik, J., 1886. Ueber Elliptera omissa Egg. - Wiener Entomologische Zeitung 5: 337-352. Mik, J., 1894. Ein Beitrag zur Biologie einiger Dipteren. 1. Die Metamorphose von Dactylolabis denticulata Bergr. - Wiener Entomologische Zeitung 13: 261- 270. Miiggenberg, F. H., 1901. Larve und Puppe von Cylindro- toma glabrata (Meigen) 1818, ein Beitrag zur Kennt- nis der Tipuliden. - Archiv für Naturgeschichte 67 (Beiheft): 114-136. Needham, J. G., 1903. Some new life histories of Diptera. — New York State Museum Bulletin 68: 279-287. Needham, J. G., 1908. Report of the entomological field station conducted at Old Forge, N.Y., in the summer of 1905. - New York State Museum Bulletin 124: 156- 248 + 22 pl. Needham, J. G. & R. O. Christensen, 1927. Economic insects in some streams of Northern Utah. - Utah Agricultural Experimental Station Bulletin 201: 1-36. Nielsen, P., O. Ringdahl & S. T. Tuxen, 1954. Diptera, 1 (exclusive of Ceratopogonidae and Chironomidae). — Zoology of Iceland 3 (48A): 1-189. Nowell, W. R., 1951. The Dipterous family Dixidae in western North America (Insecta: Diptera). -Microen- tomology 16: 187-270. 264 Nowicki, M., 1867. Beschreibung neuer Dipteren. - Ver- handlungen des Zoologischen-Botanischen Gesell- schaft in Wien 17: 337-354. Okely, E. F, 1979. The larval morphology of Erioptera trivialis Meigen and Molophilus occultus de Meijere, with additional notes on Tricyphona immaculata Mei- gen (Diptera, Tipulidae: Limoniinae). - Proceedings of the Royal Irish Academy 79 (B): 235-246. Oldham, J. N., 1926. On the larval stage of Pedicia rivosa L. - Proceedings of the Royal Physical Society of Edinburgh 21: 33-64. Oosterbroek, P., 1986. A phylogenetic classification of the Tipuloidea, based on pre-imaginal characters. - Ab- stracts of the First International Congress on Dipte- rology, Budapest, August 1986: 178. Oosterbroek, P., 1989. Tipulidae. - In: Catalog of the Diptera of the Australasian and Oceanian Regions (edited by N. L. Evenhuis): 53-166. - Bishop Museum Press, Honolulu; Brill, Leiden. Osten Sacken, R., 1869. Monographs of the Diptera of North America, 4. - Smithsonian Miscellaneous Col- lections 219: 1-345. Pastejrik, J., 1908. Metamorphosa nekterych dipter. - Casopis Ceske Spolecnosti Entomologicke 5: 27-31 (in Czech). Perraudin, J., 1961. Recherches sur l’anatomie cephalique des larves de Bibionides et de Lycoriides (Dipt. Nematocera). - Travaux du Laboratoire de Zoologie et de la Station Aquicole Grimaldi de la Faculté des Sciences de Dijon 41: 1-47. Perris, M. E., 1847. Notes pour servir a l’histoire des metamorphoses de diverses espèces de dipteres. (Deuxième partie) (1). - Annales de la Société Ento- mologique 1847: 331-351. Peterson, A., 1960. Larvae of Insects, Vol. 2 (4 ed.). —- Columbus, Ohio: 416 pp. Peterson, A., 1965. Larvae of Insects, Vol. 1 (5 ed.). - Columbus, Ohio: 315 pp. Peterson, B. V., 1981. Simuliidae. — In: Manual of Nearctic Diptera, Vol. 1. (edited by J. F McAlpine et al.): 355- 391, Biosystematic Research Institute, Ottawa, Onta- rio, Monograph 27: i-vi, 1-674. Peus, F, 1952. 17. Cylindrotomidae. — Fliegen der Pa- laearktischen Region, 3(5)3, Lief. 169: 1-80. Picado, C., 1913. Les broméliacées épiphytes. - Bulletin des Sciences de France et Belgique, 47: 215-360. Pierre, C., 1924. Tipulidae nouveaux. - Encyclopédie En- tomologique, B II, Diptera 1: 79-93. Pierre, C., 1926. Tipulidae nouveaux pour la faune de France. - Encyclopédie Entomologique, B II, Diptera 3:921222. Pilgrim, R. L. C., 1972. The aquatic larva and the pupa of Choristella philpotti Tillyard, 1917 (Mecoptera: Nan- nochoristidae). — Pacific Insects 14: 151-168. Plachter, H. 1979. Zur Kenntnis der Präimaginalstadien der Pilzmücken (Diptera, Mycetophiloidea). - Zoolo- gische Jahrbücher, Abteilung für Anatomie und On- togonie der Tiere 101: 168-266, 271-392, 427-455. Poisson, R., 1932. Sur la biologie de Geranomyia unicolor Haliday. - Societé Entomologique de France (Livre de Centenaire): 311-318. Pokorny, P., 1978. Zur Morphologie und Taxonomie der Eloeophila-Larven und -Puppen (Diptera, Limonii- dae). - Acta Entomologica Bohemoslovaca 75: 130- 142. Quate, L. W. & J. R. Vockeroth, 1981. Psychodidae. - In: Manual of Nearctic Diptera, Vol. 1. (edited by J. F. McAlpine et al.): 293-300, Biosystematic Research Institute, Ottawa, Ontario, Monograph 27: i-vi, 1-674. Reusch, H., 1988. Untersuchungen zur Faunistik, Phä- nologie und Morphologie der Limoniidae im Nieder- sächsischen Tiefland (Insecta, Diptera, Nematocera). - Dissertation Universität Hamburg: 154 pp, 77 pl. Ring, R. A., 1978. Spiracular gills of the pupa of the intertidal crane fly, Limonia (Idioglochina) marmo- rata (Diptera: Tipulidae). - Canadian Entomologist 110: 1277-1280. Röder, G., 1984. Morphologische Untersuchungen an Praetarsen von Diptera und Mecoptera (Insecta). - Dissertation Friedrich Alexander Universität Erlan- gen-Nürnberg: 393 pp. Rogers, J. S., 1926. Notes on the biology and immature stages of Gonomyia (Leiponeura) pleuralis (Will.), Tipulidae, Diptera. - Florida Entomologist 10: 33-38. Rogers, J. S., 1927a. Notes on the biology of Atarba picticornis Osten Sacken. Tipulidae-Diptera. - Florida Entomologist 10: 49-55. Rogers, J. S., 1927b. The immature stages of Gnophomyta jacobsoni Alexander (Dipt.). - Supplementa Entomo- logica 16: 77-80. Rogers, J. S., 1927c. Notes on the life history, distribution and ecology ot Diotrepha mirabilis Osten Sacken. — Annals of the Entomological Society of America 20: 23255} Rogers, J. S., 1927d. Notes on the biology and immature stages of Geranomyia (Tipulidae, Diptera). 1. Gerano- myia rostrata (Say).- Florida Entomologist 11: 17-26. Rogers, J. S., 1928. Notes on the biology of Gnophomyia luctuosa Osten Sacken, with descriptions of the imma- ture stages. - Annals of the Entomological Society of America 21: 398-406. Rogers, J. S., 1932. On the biology of Limonia (Dicranomyia) floridana (Osten Sacken). — Florida Entomologist 15: 65-70. Rogers, J. S., 1933. Contribution toward a knowledge of the natural history and immature stages of the crane flies. 1. The genus Polymera Wiedemann. — Occa- sional Papers of the Museum of Zoology, University of Michigan 268: 1-13. Rogers, J. S. & G. W. Byers, 1956. The ecological distri- bution, life history, and immature stages of Lipsothrix sylvia. - Occasional Papers of the Museum of Zoology, University of Michigan, Ann Arbor 572: 1-14. Rohdendorf, B., 1964. The historical development of the Diptera. — Trudy Paleontologicheskogo Instituta, Akademiya Nauk SSR 116: 1-92. (in Russian; English translation: University of Alberta Press, Alberta, 1974). Rozkosny, R. 1980. Diptera, Tipulidae and Cylindrotomi- dae. - In: Keys to the waterlarvae of insects (edited by R. Rozkosny): 235-244. Academia, Praha. Rozkosny, R. & P. Pokorny, 1980. Limoniidae. - In: Keys to the waterlarvae of insects (edited by R. Rozkosny): 245-257. Academia, Praha. Saether, O. A. 1971. Notes on general morphology and terminology of the Chironomidae (Diptera). - Cana- dian Entomologist 103: 1237-1260. Satchell, G. H., 1953. On the early stages of Bruchomyia argentina Alexander (Diptera: Psychodidae). — Pro- ceedings of the Royal Entomological Society of Lon- don (A) 28: 1- 12. OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Saunders, L. G., 1928. Some marine insects of the Pacific coast of Canada. - Annals of the Entomological So- ciety of America 21: 521-545. Saunders, L. G., 1930. The early stages of Geranomyia unicolor Haliday, a marine tipulid. - Entomologist’s Monthly Magazine 16: 185-187. Savchenko, E. N., 1966. Tipulidae. - Fauna Ukrainy 14(1): 1-551 (in Ukranian). Savchenko, E. N., 1979. Phylogenie und Systematik der Tipulidae. - Tijdschrift voor Entomologie 122: 91- 126. Savchenko, E. N., 1982. Limoniidae: Eriopterinae. - Fauna Ukrainy 14(3): 1-335 (in Ukranian). Savchenko, E. N., 1983a. The Limoniid flies of South Primorye. - Nauk Dumka, Kiev: 156 pp (in Russian). Savchenko, E. N., 1983b. Crane-flies (Fam. Tipulidae), Introduction, Subfam. Dolichopezinae, subfam. Tipu- linae (start). - Fauna USSR, Diptera, 2(1-2), (N.S.) 127: 1-585 (in Russian). Savchenko, E. N., 1985. Limoniidae: Limoniinae. Fauna Ukraini 14(4): 1-180 (in Russian). Savchenko, E. N., 1986. Limoniidae: Introduction, Pedi- ciinae, Hexatominae. — Fauna Ukraini 14(2): 1-379 (in Russian). Savchenko, E. N., 1989. Limoniidae fauna of the U.S.S.R. — Akademia Nauk Ukranian SSR., Kiev: 377 pp (in Russian). Savchenko, E. N. & G. O. Krivolutskaja, 1976. Limoniidae of the south Kuril and south Sakhalin. - Akademia Nauk Ukranian SSR., Kiev: 160 pp (In Russian). Savchenko, E. N., P. Oosterbroek & J. Stary, 1992. Limo- niidae. — Catalogue of Palaearctic Diptera, Vol. 1 (Edited by A. Sods & L. Papp) (in press). Seguy, E., 1942. Un nouveau Diptere Molophilus pauliani nov. sp. sur le littoral de la Manche. - Bulletin du Laboratoire de Dinard 24: 77-80. Seligo, A., 1931. Zur Kenntnis der Bodentierwelt des Mariensees. — Bericht des Westpreussischen Botan- ish-Zoologischen Vereins 53(9): 1-33. Seurat, L.-G., 1924. Moeurs et évolution d’un Tipulide a larve et nymphe marines. - Bulletin de la Sociéte d'Histoire Naturelle de l'Afrique 15: 113-121. Sinclair, B. J., 1988. The madicolous Tipulidae (Diptera) of eastern North America, with descriptions of the biology and immature stages of Dactylolabis montana (Osten Sacken) and D. hudsonica Alexander (Diptera: Tipulidae). - Canadian Entomologist 120: 569-573. Smith, K. G. V., 1989. An introduction to the immature stages of British flies. - Handbooks for the Identifica- tion of British Insects 10(14): 1-280. Snodgrass, R. E., 1960. Facts and theories concerning the insect head. - Smithsonian Miscellaneous Collections 142: 1-61. Spencer, G. J., 1930. Notes on Phalacrocera species, an aquatic crane fly (Diptera, Tipulidae). - Proceedings of the Entomological Society of British Columbia 27: 15-16. Stannius, H., 1829. Zur Verwandlungsgeschichte der Limnobia xanthoptera Meigen. — Beiträge zur Ento- mologie, Breslau 1: 202-206. Steiner, P., 1930. Studien an Panorpa communis. — Zeitschrift fiir Morphologie und Okologie der Tiere 17: 1-25, 26-67. Studnitz, G. von, 1932. Die statische Funktion der sog. ‘pelotaktischen’ Organe (‘Schlammsinnesorgane’) der Limnobiidenlarven. - Zoologische Jahrbiicher, Abtei- 265 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 lung für Allgemeine Zoologie 50: 419-446. Swezey, O. H., 1915. A leaf-mining cranefly in Hawaii. — Proceedings of the Hawaiian Entomological Society 3: 87-89. Szadziewski, R., 1979. The immature stages of two halo- biont Diptera, Dicranomyia sera (Walker) (Limonii- dae) and Parascatopse litorea (Edwards) (Scatopsi- dae). - Polskie Pismo Entomologiczne 49: 385-388. Teskey, H. J., 1976. Diptera larvae associated with trees in North America. - Memoirs of the Entomological Society of Canada 100: 1-53. Teskey, H. J., 1981. Morphology and Terminology: Lar- vae. - In: Manual of Nearctic Diptera, Vol. 1. (edited by J. F McAlpine et al.): 65-88, Biosystematic Re- search Institute, Ottawa, Ontario, Monograph 27: i-vi, 1-674. Theowald, Br., 1957. Die Entwicklungsstadien der Tipu- liden, insbesondere der west-palaearktischen Arten. — Tijdschrift voor Entomologie 100: 195-308. Theowald, Br., 1967. Familie Tipulidae, Larven und Puppen. - Bestimmungsbücher zur Bodenfauna Euro- pas 7: 1-100. Tokunaga, M., 1930. The morphological and biological studies on a new marine cranefly, Limonia (Dicranomyia) monostromia, from Japan. -Memoirs of the College of Agriculture, Kyoto Imperial Univer- sity 10: 1-93. Tokunaga, M., 1933. A marine crane fly, Limonia (Dicranomyia) trifilamentosa, of the pacific coast of Japan, with special reference to the ecology and the morphology of its immature stages. - Philippine Jour- nal of Science 50: 327-344. Tokunaga, M., 1936. Three marine crane-flies from Ja- pan. - Annotationes Zoologicae Japonensis 15: 460- 468. Tokunaga, M., 1939. Female and larva of the marine crane fly Limonia (Idioglochina) gloriosa Tokunaga. - An- notationes Zoologicae Japonensis 18: 101-104. Tokunaga, M., H. Ishida & A. Nobushi, 1954. Crane flies injurous to useful mushrooms in Japan, with revision of the Japanese Ula species and descriptions of a known and three new species and immature stages of two species. - Science Reports of the Sayko University of Agriculture, Kyoto 6: 1-10. (Not consulted, cited from Lindner, 1959.) Tonnoir, A. L., 1926. A new sub-apterous and sub-halter- less crane fly. - Records of the Canterbury Museum 3: 27-30. Toye, S. A., 1967. The larva of the marine Tipulid, Ge- ranomyia unicolor Haliday. - Proceedings of the Royal Entomological Society of London, (A) 42: 167- 170. Ussing, H. J., 1929. Dicranomyia didyma Meigen. - Flora og Fauna 1929: 81-84. Vaillant, FE, 1951. Sur Orimarga hygropetrica n.sp. (Dip- tere, Limnobiidae, Heliini). - Travaux du Laboratoire d’Hydrobiologie et Pisciculture de Grenoble 41-42: 43-47. Vaillant, F., 1953. Larve et nymphe de Trimicra hirsutipes Macquart (Diptera Limnobiidae Eriopterini). - Mis- sion Scientifique au Tassili des Ajjer (1949). 1 (Re- cherches Zoologiques et Médicales): 2-8. Vaillant, F, 1956. Recherches sur la faune madicole (Hy- gropétrique s.l.) de France, de Corse et d'Afrique du Nord. - Mémoires du Museum National d'Histoire Naturelle, N.S. (A) Zoologie 11: 1-252. 266 Vimmer, A., 1906. Uber die Mundwerkzeuge der Tipu- linen und Pachyrhinen-larven. - Casopis Ceské Spo- lecnosti Entomologické 3: 37-49 (in Czech and Ger- man). Vimmer, A., 1912. Ueber den Hypopharynx einiger Dip- terenlarven aus der Unterordnung Orthorrapha. - Societas Entomologica 27(23): 103-103. Vimmer, A., 1924. Larve und Puppe der Gruppe Poly- neura (Dipt.). - Sbornik Entomologického Oddeleni Narodniho Musea Praze 2: 133-136. Vimmer, A., 1925. Larvy a kukly dvojkridleho hmyzu stredoevropskeho. - Nakladem Ceske Graficke Unia, Praze: 349 pp, 59 pl. (in Czech). Vimmer, A., 1928. Eine interessante, in Boehmen gefun- dene amerikanische Dipteren-Larve (Dipt., Limnobii- dae). - Sbornik Entomologického Oddeleni Närod- niho Musea Praze 6: 67-71. Vimmer, A., 1929. Übersicht des Tracheensystems und der Trachealisation der innere Organen der Dipteren- larven. - Casopis Ceské Spolecnosti Entomologické 11: 68-88. Walton, G. A., 1944. An undescribed aquatic tipulid (Dipt.) larva. - Journal of the Society for British En- tomology 2: 166-168. Wardle, R. A., 1926. 2. The respiratory system of con- trasting types of cranefly larvae. - Proceedings of the Zoological Society of London 1926: 25-48. Wardle, R.A. & E.A. Taylor, 1926. 1. The cephalic ske- leton of contrasting types of cranefly larvae. - Pro- ceedings of the Zoological Society of London 1926: 1- 24. Watrous, L. E. & Q. D. Wheeler, 1981. The outgroup comparison method of character analysis. -Systematic Zoology 30: 1-11. Wesenberg-Lund, C., 1943. Biologie der Süßwasserinsek- ten. - Springer, Berlin: 10 + 682 pp. Wiedenska, J., 1987. Morphologie der Larven und Puppen einiger Phylidorea-Arten (Diptera, Limonii- dae). - Tijdschrift voor Entomologie 130: 33-47. Wiley, E. O., 1981. Phylogenetics. The theory and practice of phylogenetic systematics. - J. Wiley & Sons, New York: 16 + 439 pp. Williams, F X., 1943. Biological studies in Hawaiian water-loving insects. 3. Diptera or Flies. C. Tipulidae and Psychodidae. - Proceedings of the Hawaiian En- tomological Society 11: 313-338. Willmann, R., 1989. Evolution und phylogenetisches Sys- tem der Mecoptera (Insecta: Holometabola). — Ab- handlungen der Senckenbergischen Naturfor- schenden Gesellschaft 544: 1-153. Winterbourn, M. J. & K. L. D. Gregson, 1989. Guide to the aquatic insects of New Zealand. - Bulletin of the Entomological Society of New Zealand 9: 1-95. Wirth, W. W. & A. Stone, 1956. Aquatic Diptera. Family Tipulidae. - In: Aquatic Insects of California (edited by Usinger): 377-384. — University of California Press, Berkeley. Wolff, B., 1922. Schlammsinnesorgane (pelotaktische Or- gane) bei Limnobiinenlarven. - Jenaische Zeitschrift fiir Naturwissenschaft 58: 77-144. Wood, D. M. & A. Borkent, 1989. Phylogeny and classi- fication of the Nematocera. — In: Manual of Nearctic Diptera, Vol. 3. (edited by J. F McAlpine et al.): 1333- 1370, Biosystematic Research Centre, Ottawa, Onta- rio, Monograph 32: i-vi, 1333-1581. OOSTERBROEK & THEOWALD: Phylogeny of Tipuloidea Wood, H. G., 1952. The crane-flies of the South-West Received: 28 June 1991 Cape (Diptera, Tipuloidea). - Annals of the South Accepted: 7 October 1991 African Museum 39: 1-327. Young, C. W., 1987. A revision of the crane fly genus Dicranoptycha in North Americ:. Kansas Univer- sity Science Bulletin 53: 215-274. 267 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 BOOK REVIEW Moller Pillot, H. K. M. & R. E M. Buskens, 1990. De larven der Nederlandse Chironomidae (Diptera). Deel C: Autoekologie en verspreiding. [The larvae of the Dutch Chironomidae (Diptera). Part C: Autecology and distribution]-Nederlandse Faunistische Medede- lingen 1C: 1-87, figs 1-2, tables 1-5, maps 1-182. [Stichting European Invertebrate Survey-Nederland; send orders to ‘The Library, Nationaal Natuurhisto- risch Museum, P.O. Box 9517, 2300 RA Leiden, Ne- derland’. Price 20 Dutch Guilders. Price of parts 1A- C together 100 Dutch Guilders. Text in Dutch, but an explanation in English of the table containing all ecological information is available on request]. Keys for the identification of larvae of Dutch Chironomidae by H. Moller Pillot were published in the series Nederlandse Faunistische Mededelin- gen (parts 1A and 1B) in 1978 and 1984. In both works keys are given for the majority of genera, but the identification of species appeared to be difficult or impossible in many cases. For this reason Moller Pillot introduced a system of species-groups, aggre- gates, and sometimes even artificial names, e.g. ‘Chaetocladius spec. Herkenbosch’, for larvae that could clearly be recognised although no unequivocal correlation could be made with species already des- cribed as adults. Thus a system of about 200 larval taxa was created, covering all Chironomidae, except for the Tanytarsini. Chironomidae have proved to be good indicators to various conditions of the aquatic and semi-aquatic environment. For this reason they play an impor- tant role in several systems for the biological assess- ment of water quality. As these systems came into use on a wide scale throughout The Netherlands during the eighties, the keys by Moller Pillot were widely used soon after publication. New water qual- ity systems were developed as well, in which the taxa of Moller Pillot were incorporated. Already from the beginning of his studies on Chironomidae, Moller Pillot collected ecological data from his own investigations as well as data coming forth from the use of the keys in combina- tion to large environmental research projects and the systems for the biological assessment of water quality. Dozens of students from all over The Ne- therlands put their data to disposal. Based on these data, Moller Pillot and Buskens have now prepared the third and final part (part 1C) of the work on Dutch Chironomidae. The book deals with the autecology and the geo- graphic distribution of theca. 200 taxa distinguished in parts 1A and 1B. The nomenclature from parts 1A and 1B has been maintained as muchas possible; a brief list on page 78 in part 1C indicates the names that had to be changed or were added since. 268 The autecology is summarized in a brief but elegant way. In a large table, which represents the most important part of the book, the ecological response of all taxa to a number of environmental factors is given. Factors treated in the table are water type, habitat, oxygen, chloride concentration and the acidity in combination with the trophic level. Each factor is split up in a number of conditions. The ecological response of each taxon for each condition of each factor is roughly indicated on a scale, varying between 0 and 4. These numbers indicate the rel- ative abundance of the taxa under the conditions mentioned, in a range from total absence to a high abundance. The authors emphasize that habitat preference of species is often not determined by the mean values of environmental factors, but by critical values instead. For instance, four conditions of the factor oxygen are given in the table: stable (satu- ration always more than 50%), instable (saturation level between 10 and 50% at least once a day), saturation level below 5% for at most a few hours a day, and saturation level below 5 % for many hours each day in summer. This approach offers more adequate ecological information than a given re- sponse to mean oxygen values would do. Additional information for each taxon is given in a series of short notes, concerning the life cycle and ecological details not mentioned in the large table. In a number of cases the distribution patterns of taxa are discussed from an ecological point of view. The distribution patterns in The Netherlands are represented by 182 maps, using the 10 km UTM- grid. Only records checked by the authors were included in these maps. For a better understanding of the distribution patterns maps of the distribution of soil types and brackish water in The Netherlands were added. In addition to the maps, the species abundance is mentioned in the large table as well, giving the number of 10 km squares from the maps and by using a code for the number of 5 km squares from the Dutch National Grid. The parts A, B and C together reflect the results of many years of investigations and all data pres- ented are original. This is not only important from a scientific point of view. Especially the suitability for practical use, of which an exampie is worked out in part C, makes the work most valuable. Although the text is in Dutch, the books are of importance for chironomid workers outside The Netherlands as well. The keys (parts A and B) contain figures of relevant parts of the body of all taxa included, while the data on the ecology and distribution (in part C) are readily accessible by the concise way of presentation. [A. W. M. Mol] GARY M. STONEDAHL International Institute of Entomology, London REVIEW OF THE ORIENTAL GENUS ANGERIANUS DISTANT (HETEROPTERA: MIRIDAE) Stonedahl, G. M., 1991. Review of the Oriental genus Angerianus Distant (Heteroptera: Miridae). - Tijdschrift voor Entomologie 134: 269-277, figs. 1-20. [ISSN 0040-7496]. Published 18 December 1991. The genus Angerianus Distant is diagnosed and redescribed, and its recent placement in the subfamily Deraeocorinae is confirmed. The new species A. mindanao and A. palawa- nensis from the Philippine Islands, and A. pallidus from Sabah and Sarawak are diagnosed and described. Lectotypes are designated for the Distant species A. fractus and A. maurus based on syntypic material preserved in the Natural History Museum, London. Illustra- tions are provided for the male genitalic structures of all species, and scanning electron micrographs are presented for the scent gland ostiole and pretarsus of A. maurus. A key is provided which uses external features and characters of the male genitalia to distinguish the five known species. G. M. Stonedahl, International Institute of Entomology, 56 Queen's Gate, London SW7 SJR, U. K. Key words. - Heteroptera; Miridae; Deraeocorinae; Angerianus, review; new species; Oriental region. Angerianus was described by Distant (1904) to accommodate two new species, fractus and maurus, collected at Myitta in the Tenasserim Valley, Burma. The genus was placed in the tribe Dicy- phini by Carvalho (1952), but Cassis (1984), in a revision of this group, moved Angerianus into the Deraeocorinae. Stonedahl and Cassis (1991) pres- ent character information suggesting that Angeri- anus is most closely related to the Old World genus Fingulus Distant. In conjunction with that study, three new species of Angerianus were discovered in the entomological collections of the Bernice P. Bishop Museum, Honolulu. Descriptions of these species are provided here along with redescriptions of fractus and maurus, and a key to distinguish the five known species. All measurements are given in millimeters. Body length is measured from the apex of the tylus to the deepest point of the cuneal incisure. Abbre- viations used in the locality data to denote speci- men depositories correspond to the institutions listed in the acknowledgments. SYSTEMATIC ACCOUNT Angerianus Distant Angerianus Distant, 1904: 437-438. - Type species: An- gerianus fractus Distant. Angerianus: Reuter 1910: 161 (cat.); Carvalho 1952: 72 (classif., cat.), 1955: 60 (key), 1957: 182 (cat.); Cassis 1984: 166 (diag., subfam. placement). Diagnosis. - Recognized by the head not pro- duced anteriad of eyes, with vertical frons and tylus; well developed cephalic neck; long first antennal segment; broad pronotal collar; lateral margins of pronotum constricted at level of anterior and pos- terior borders of calli; strongly deflexed hemelytra with broad, deep cuneal incisure; long, tapered femora (fig. 1); strongly protruding scent gland ostiole (figs. 2, 3); and structure of the male geni- talia, especially the vesica without lobal sclerites, and no serrate plates bordering secondary gono- pore. Redescription of male. - Macropterous, length 2.19-2.77; pale brownish yellow to dark brown general coloration; dorsal surface shiny, smooth; collar and disc of pronotum finely to coarsely punc- tate; dorsal vestiture with moderate to dense dis- tribution of long, pale setae, or sometimes nearly glabrous with shorter, scattered setae. Head: Ver- tical anteriorly, not produced distad of eyes, with weakly produced tylus; broader than long in dorsal view with vertex nearly twice as broad as dorsal width of eye; tapered posteriad of eyes into distinct necklike region, with eyes well removed from an- terior margin of pronotum; neck anteriorly with 269 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Fig. 1. Angerianus maurus, dorsal habitus female. > transverse dorsal depression; eyes occupying nearly entire height of head in lateral view. Anten- nae: Long, thin, inserted well dorsad of median level of eyes; length of segment I equal to or greater than width of head across eyes; segments II-IV with moderately long, suberect, pale setae. Labium: Reaching to meso- or metacoxae; segments I-III similar in length, segment IV slightly longer. Pron- otum: Strongly narrowed anteriorly and noticeably constricted at level of posterior margin of calli and at juncture of collar and calli; collar much broader than diameter of antennal segment I, flattened or weakly convex dorsally; calli confluent medially, 270 reaching lateral margins of pronotum, surface smooth or faintly rugulose, sometimes with long, pale setae; posterior margin of disc broadly convex, concealing most of mesoscutum and sometimes anterior margin of scutellum; scutellum elevated above resting hemelytra; metathoracic scent effer- ent system with strongly protruding ostiole, eva- porative area not or only slightly extending dorsad of upper margin of ostiole. Hemelytra: Strongly deflexed at cuneal fracture; lateral margins nearly straight anteriorly, curving inward distally to broad, deep cuneal incisure; costal vein and claval suture marked by row of coarse punctures; cuneus relatively long, with curved outer margin; mem- brane reaching well beyond apex of abdomen, with large primary cell and obsolete secondary cell. Legs: Femora elongate, tapered, broadest basally; tibiae and tarsi cylindrical, with pale, suberect setae; pre- tarsal claws strongly curved with prominent basal tooth (fig. 5). Genitalia: Genital capsule: Small, narrow, ventral region between paramere sockets slightly produced posteriorly; aperture large, pos- teriorly directed; paramere sockets asymmetrical with left socket slightly larger and ventrad of right socket. Left paramere: L-shaped with weakly to moderately produced sensory lobe; shaft long, ta- pered distally to blunt, sometimes truncate apex. Right paramere: Small, ovate to lanceolate, some- times strongly narrowed distally; apex blunt. Phal- lotheca: Short, conical, narrowly opened dorsodis- tally. Vesica: Composed of multilobed membra- nous sac surrounding distal region of ductus seminis, and simple, unmodified secondary gono- pore opening into depressed area between mem- branous lobes. Lobes of vesica sometimes with patches of blunt tubercles, but lacking lobal scler- ites. Basal region of vesica not surrounded by sclerotized skirt, and without serrate plates border- ing secondary gonopore. Female. - Macropterous, length 2.22-2.73; sim- ilar to male in general structure and coloration except as noted in species descriptions. Genitalia not examined. Distribution. - Tropical Asia from Nepal east to Taiwan, the Philippine Islands and Sabah, and south to Java. Discussion. - Distant (1904) placed Angerianus in the Division Cylaparia (Cylapinae), but Reuter (1910), uncertain of its relationships, treated this genus as incertae sedis in his classification of the Miridae. Apparently unaware of its distinct de- raeocorine pretarsus and male genitalia, Carvalho (1952) moved Angerianus into the tribe Dicyphini (Phylinae). The correct subfamilial placement of Angerianus was established by Cassis (1984), who noted that members of this genus have a typical deraeocorine pretarsus, with cleft claw bases and setiform parempodia. Cassis also observed that the structure of the mesepimeric spiracle and the me- taepisternal scent efferent system were like those of the Deraeocorinae and not the Dicyphini. The observations of Cassis are confirmed in the present study, and it is further noted that the male genitalic structures of Angerianus support its placement in the Deraeocorinae, particularly the shape of the left paramere and the depressed, unmodified se- condary gonopore of the vesica. The relationship of Angerianus to other de- raeocorine genera is not well established. Cassis STONEDAHL: Review of Angerianus (1984) tentatively regarded Angerianus as most closely related to the Hyaliodini, based primarily on the strongly protruding, tubelike auricle of the metaepisternal scent efferent system. Comment- ing on the similarities between Angertanus and Stethoconus Flor, Cassis questioned the placement of the later genus in the Clivinemini by Kerzhner (1967). Stethoconus has since been moved into the Hyaliodini by Henry et al. (1986). In a recent revision of the Old World deraeoco- rine genus Fingulus Distant, Stonedahl and Cassis (1991, in press) suggested a possible sister group relationship between this genus and Angerianus based on characters of the external morphology and male genitalia. Further, these authors noted that Fingulus and Stethoconus, while superficially similar, have very different male genitalia and head morphology, indicating that they are not as closely related as originally suggested by Cassis (1984). Based on these preliminary findings, I consider Angerianus to be most closely related to Fingulus, but can say little about the relationships of these genera to other Deraeocorinae, except to suggest a possible link with the Hyaliodini. Key to species of Angerianus Collar and disc of pronotum with numerous coarse punctures and long, pale setae ....... 2 — Collar and disc of pronotum with scattered fine punctures and short setae, or impunctate and appearimetglabrous u. TETTE 4 2. Antennal segment I narrowly pale basally, brown or black distally; length of segment no- ticeably greater than width of head across eyes; male genitalia as in figures 12-14 .............. Re UU ee 0002 mindanao sp. n. — Antennal segment I darkened basally and pale distally; length of segment about equal to width or headiaGrossieyesie eee ee ere ee eer rere 3 3. Fore femora mostly pale, sometimes lightly infuscated medially or rarely more extensively darkened distally; scutellum usually with large, pale spot distomedially; length of antennal segment II 1.24-1.44; male genitalia as in fig- URES 028: ie OR fractus Distant — Fore femora mostly fuscous, basal fourth pale; scutellum uniformly darkened, without pale spot medially; length of antennal segment II 1.02-1.17; male genitalia as in figures 9-11 DIRE oc AIA SUSA Re maurus Distant 4. Pale brownish yellow general coloration; head uniformly pale; length of labium about equal to posterior width of pronotum; male genitalia as ID IQUTES PIS 2 RER pallidus sp. n. - Dark brown general coloration; head infus- cated posteriad of eyes, and with dark stripe reaching from dorsal margin of antennal fos- 737/31 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 sae to posterior margin of head; length of la- bium much greater than posterior width of pronotum; male genitalia as in figures 15-17 LOSE ATTIRE MAS | palawanensis sp. n. Angerianus fractus Distant (figs. 6-8) Angerianus fractus Distant, 1904: 438, fig. 283. Lectotype Q (new designation): Label 1, ‘Type, H.T.’ [red-bor- dered disc]; 2, ‘Angerianus fractus Dist.’ [handwritten]; 3, ‘Tenass Vall, Myitta, (Doherty).’; 4, ‘Distant Coll., 1911-383’; 5 (red label added here), ‘Lectotype, Angerianus fractus Distant, det. by G. M. Stonedahl, 1991.’ (NHML) [examined]. Angerianus fractus; Carvalho 1957: 182 (cat.). Figs. 2-5. Scanning electron micrographs of Angerianus maurus. - 2, Lateral view of thorax; 3, Ostiole and evaporative Diagnosis. - Recognized by the coarsely punc- tate collar and disc of the pronotum; bicolored first antennal segment, with distal region pale; dark scutellum, usually with large, pale spot distome- dially; and by the structure of the male genitalia (figs. 6-8). This species is further distinguished from maurus by the longer second antennal seg- ment and mostly pale fore femora. Description of male. - Length 2.37-2.52; yellow- ish brown to dark brown general coloration. Head: Width across eyes 0.66-0.70, vertex 0.30-0.33; mostly pale anteriorly and dorsally; gula and neck- like region posteriad of eyes brown or dark brown; lorum, buccula and gena usually lightly to heavily area of metathoracic scent efferent system; 4, Metafemur showing trichobothria; 5, Pretarsus. 22 tinged with fuscous; vertex with V-shaped mark running from antennal fossae to depressed line marking anterior margin of neck. Antennae: I, length 0.62-0.70, basal half to two-thirds brown or dark brown, or in paler specimens sometimes only narrowly darkened basally, distal region pale yel- low or dirty white; II, length 1.25-1.44, pale brow- nish yellow; III & IV, brown. Labium: Length 1.17- 1.22; reaching between mesocoxae; segments I-III reddish brown, segment IV brownish yellow. Pron- otum: Posterior width 1.16-1.25; collar and disc coarsely punctate; yellowish brown to dark brown, lighter specimens usually considerable darker lat- erally; scutellum brown or dark brown, usually with large pale spot distomedially; scent gland ostiole grayish white, much paler than surrounding pleura. Hemelytra: Dark yellowish brown to mostly dark brown - lighter specimens usually with apex of clavus, distal region of embolium and cuneus darker brown or brownish red; darker spec- imens with apical angle of corium and base of cuneus noticeably paler; membrane lightly to mod- erately tinged with fuscous basally, sometimes to level of apex of cuneus, veins brown or dark brown. Legs: Grayish yellow or brownish yellow; femora of lighter specimens usually with middle third in- fuscated, darker specimens usually with femora more extensively darkened distally and with base of tibiae narrowly darkened. Genitalia: Left paramere and vesica as in figures 6-8. Female. - Length 2.34-2.50; width across eyes 0.65-0.67, vertex 0.31-0.32; length of antennal seg- ment I 0.62-0.67, segment II 1.24-1.29; length of labium 1.08-1.20; posterior width of pronotum 1.17-1.27. Similar to male in structure and color, except second antennal segment usually much shorter, and pronotum broadly pale behind calli and with distinct pale stripe medially on disc. Distribution. - Burma, Laos, Nepal, Thailand and Vietnam. Discussion. — This species is quite variable in general coloration, ranging from yellowish brown with limited darker brown markings to mostly dark brown (see description of male). Females are less variable than males in coloration, and are always easily distinguished from other species of Angert- anus by the pale, median stripe on the pronotal disc. The original description of A. fractus gives Myitta, Tenasserim as the type locality. Although there is no indication of the number and sex of the specimens that Distant examined, it is clear from the habitus illustration and text that he was des- cribing the female of this species. Two specimens of A. fractus from the type locality are deposited in the Natural History Museum in London. The fe- male is here designated the lectotype, but it is STONEDAHL: Review of Angerianus Figs. 6-8. Male genitalia of Angerianus fractus. - 6, Ves- ica; 7, Left paramere, dorsal view; 8, Left paramere, pos- terolateral view. questionable whether the male specimen then be- comes a paralectotype, as there is no evidence to suggest that Distant ever looked at specimens of this sex. The head and prothorax of the lectotype have at some point been dislodged from the body and are now glued to the card point. All of the legs are missing, and only the first segment and base of the second segment of the right antenna remain. Specimens examined. - Burma: Tenasserim: 1 4, same data as lectotype (NHML); 1 ® Dawna Hills, 2000-3000 ft, March 2-3, 1908 (NHML, Distant Coll.). - Laos: Vientiane Prov.: 1 9, Ban Van Eue, SE of Phou Kow Kuei, 800 m, April 14, 1965, J.L. Gressitt (BISH); 1 4, Ban Van Eue, Dec. 31, 1965, native collector (BISH). - Nepal: 1 &, Chitwan, Sauraha, Dec. 27, 1981-Jan. 9, 1982, Shorea robusta ('Sal’) forest, on low vegetation, L. Jessop (NHML); 2 6, 1 Q, Mugling, 900-1500 ft, Jan. 14-Feb. 5, 1982, mixed scrub on south facing slopes, L. Jessop (AMNH, NHML). — Thai- land: Chiang Mai Prov.: 1 6, 1 9, Banna, 108 m, May 5- 10, 1958, T.C. Maa (BISH); 1 8, 1 9, Chiangdao, 450 m, April 5-11, 1958, T.C. Maa (BISH); 1 Q Doi Suthop, Nov. 14, 1957, J.L. Gressitt (BISH); 1 6, 4 9, Fang, April 12-19, 1958, T.C. Maa (BISH); 1 6, Fang Hort. Exp. Stn., 550-600 m, Oct. 20, 1981 (UZMC). Trang Prov.: 1 , Khaophap- pha, Khaochang, 200-400 m, Jan. 13, 1964, G.A. Samuel- son (BISH). - Vietnam: 1 @, 1 9, Ban Me Thout, 500 m, Dec. 20-24, 1960, C.M. Yoshimoto (BISH); 1 9, 6 kms of Dalat, 1400-1500 m, June 9-July 7, 1961, N.R. Spencer (BISH). AUD TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 9-11. Male genitalia of Angeria- nus maurus. - 9, Vesica; 10, Left para- mere, dorsal view; 11, Left paramere, posterolateral view. Angerianus maurus Distant (figs. 1-5, 9-11) Angerianus maurus Distant, 1904: 438. Lectotype & (new designation): label 1, ‘Tenass Vall, Myitta, (Doherty).’; 2, ‘Distant Coll., 1911-383’; 3 (red label added here), ‘Lectotype, Angerianus maurus Distant, det. by G. M. Stonedahl, 1991.’ (NHML) [examined]. Angerianus maurus; Carvalho, 1957: 182 (cat.). Diagnosis. - Similar to A. fractus, but distin- guished by the darker legs, uniformly darkened scutellum and shorter second antennal segment. Males are further distinguished by the narrower vertex, longer labium, and structure of the genitalia (figs. 9-11). Description of male. - Length 2.19-2.37; brown to dark brown general coloration. Head: Width across eyes 0.66-0.70, vertex 0.24-0.26; brown or dark brown; frons, tylus, jugum, lorum and gena usually paler brownish yellow; vertex with broad, pale mark bordering inner margin of each eye. Antennae: I, length 0.63-0.67, basal two-thirds dark brown, distal region pale yellow or dirty white; II, length 1.12-1.17, pale brownish yellow; III & IV, brown. Labium: Length 1.06-1.10; reach- ing between meso- or metacoxae; segments I-II dark reddish brown, segment IV brownish yellow. Pronotum: Posterior width 1.09-1.13; collar and disc coarsely punctate; uniformly brown or dark brown; scent gland ostiole yellowish white, much paler than surrounding pleura, evaporative area sometimes lightly tinged with fuscous. Hemelytra: Brown or dark brown; anterior half of embolium, apex of corium, and basal margin of cuneus noti- ceably paler; membrane lightly tinged with fuscous basally, sometimes to level of apex of cuneus, veins brown or dark brown. Legs: Brownish yellow, with distal two-thirds of femora and base of tibiae usu- ally dark brown. Genitalia: Figures 9-11. Female. - Length 2.22-2.40; width across eyes 0.62-0.66, vertex 0.26-0.28; length of antennal seg- 274 ment I 0.64-0.69, segment II 1.02-1.11; length of labium 1.06-1.13; posterior width of pronotum 1132118: Distribution. - Widely distributed in tropical Asia from Nepal, east to Taiwan and Sabah, and south to Java. Discussion. — Like A. fractus, this species was described from a unknown number of specimens collected at Myitta in the Tenasserim Valley, Burma. Four specimens apparently belonging to the original syntype series are deposited in the Natural History Museum in London. One of these is here designated the lectotype and the others become paralectotypes. Paralectotypes. - 1 & (head and pronotum miss- ing), 1 9, same data as lectotype; 1 ®, same data as lectotype except with two additional labels below specimen as follows: Label 1, ‘Type, HT. [red- bordered disc]; 2, ‘maurus Dist.’ [handwritten]. A yellow label with the following data has been added to each of these specimens: ‘Paralectotype, Ange- rianus maurus Distant, det. by G. M. Stonedahl, 1991.’ All paralectotypes are deposited in the Nat- ural History Museum, London. Specimens examined. - Cambodia: 1 9, Siem Reap (Angkor), Dec. 6, 1957, J. Szent-Ivany (BISH). - Indone- sia: Java: 1 9, S Coast, Tankubanpraku, Sept. 27, 1960, forest remnant, J.L. Gressitt (BISH). - West Malaysia: Pahang: 1 9, Dec. 22-26, 1958, T.C. Maa (BISH). Penang: 1 9, Oct. 1913, G.E. Bryant (NHML); 1 Q, Island of Penang, Baker (USNM). - Sabah: 2 6, Liawan, Jan. 14-19, 1959, sweeping, T.C. Maa (BISH); 1 9, Samawang, nr. Sandakan, July 14, 1927 (NHML). - Sarawak: 1 6, Merirai Valley, nr. Kapit, 30-300 m, July 1-6, 1958, T.C. Maa (BISH). - Nepal: 2 9, Chitwan, Sauraha, Dec. 27, 1981-Jan. 9, 1982, Shorea robusta (Sal) forest, on low vegetation, L. Jessop (NHML). — Taiwan: 1 9, Menzendo, '1-6-33,' native pi- neapple field, K. Sakimura (BISH); 1 4, Pishan, Nov. 14, 1957, T.C. Maa (BISH). — Thailand: 1 6, 3 9, Pakchong, 100 m, NE of Bangkok, Dec. 2-3, 1957, J.L. Gressitt (BISH). Chiang Mai Prov.: 1 9, Chiangdao, April 5-11, 1958, T.C. Maa (BISH); 1 @, 1 9, Fang, April 12-19, 1958, T.C. Maa (BISH). - Vietnam: 1 @, Ap Hung-Lam, 21 km NW of Dilinh, 1100 m, Sept. 29-Oct. 5, 1960, C.M. Yo- shimoto (BISH); 7 @, 2 9, Ban Me Thout; 500 m, Dec. 20- 24, 1960, C.M. Yoshimoto (BISH); 1 9, Karyu Danar, 200 m, Feb. 13-28, 1961, N.R. Spencer (BISH); 1 @, 1 9, M'Drak, E of Ban Me Thout, 400-600 m, Dec. 8-19, 1960, C. M. Yoshimoto (BISH). Angerianus mindanao sp. n. (figs. 12-14) Type material - Holotype @: Philippine Islands: Min- danao: Davao del Sur: Galog R., Mt. Apo, 6000 ft, Sept. 26, C.S. Clagg (AMNH). — Paratypes: Philippine Islands: Mindanao: Davao del Sur: 1 9 (Sept. 5) and 4 9 (Sept. 26), same data as holotype (AMNH); 1 9, Mt. Apo, Ba- traeyon, 8000 ft, Sept. 14, CS. Clagg (AMNH); 1 4, Mt. Apo, Mainit R., 6000 ft, Sept. 24, C.S. Clagg (NHML). Additional Specimens. - Philippine Islands: Minda- nao: Bukidnon: 2 9, Mt. Katanglad, 1250 m, Oct. 26, 1959, L.W. Quate (BISH); 1 & (badly damaged, with head, he- melytra and all but one leg missing), Mt. Katanglad, 1250 m, Dec. 4-9, 1959, L.W. Quate (BISH); 1 9, Mt. View Colleges, 15 km NW of Valencia, April 22-23, 1968, D.E. Hardy (BISH). Diagnosis. - Recognized by the coarsely punc- tate pronotal collar and disc, dark brown first an- tennal segment with basal sixth to one-third pale, and by the structure of the male genitalia (figs. 12- 14). This species is further distinguished from frac- tus and marus by the longer first antennal segment and darker scent gland ostiole. Description of male. - Length 2.40-2.56; brow- nish yellow to dark brown general coloration. Head: Width across eyes 0.66-0.69, vertex 0.29- 0.32; brownish yellow, usually slightly darker ven- trally, especially posteriad of eyes; head of darker specimens brown or dark brown, with frons and vertex paler brownish yellow. Antennae: I, length 0.80-0.84, fuscous, with basal sixth to one-third pale; II, length 1.15-1.19, yellowish brown; III & IV, STONEDAHL: Review of Angerianus Figs. 12-14. Male genitalia of Angerianus mindanao. — 12, Ves- ica; 13, Left paramere, dorsal view; 14, Left paramere, postero- lateral view. brown. Labium: Length 1.25-1.39; reaching be- tween meso- or metacoxae; segments I-III reddish brown, segment IV yellowish brown. Pronotum: Posterior width 1.17-1.24; collar and disc coarsely punctate; uniformly yellowish brown, sometimes with collar and calli slightly darker; darkest spec- imens with collar, calli and margins of disc dark brown and medial region of disc lighter brown; scutellum brown or yellowish brownish; scent gland ostiole brown or yellowish brown, not or only slightly paler than surrounding pleura. He- melytra: Brownish yellow to mostly dark brown; membrane sometimes tinged with fuscous basally, veins brown or brownish yellow. Legs: Uniformly brownish yellow, or in darker specimens, with dis- tal half to two-thirds of femora, and sometimes base of tibiae darker brown. Genitalia: Left para- mere and vesica as in figures 12-14. Female. - Length 2.48-2.60; width across eyes 0.64-0.66, vertex 0.31-0.33; length of antennal seg- ment I 0.77-0.84, segment II 0.99-1.06; length of labium 1.24-1.42; posterior width of pronotum 1.12-1.20. Etymology. - Named for the island of Mindanao. Distribution. - Mindanao, Philippine Islands. Discussion. - This species is quite variable in general coloration, the dorsum ranging from uni- formly pale brownish yellow (type and paratypes from Mt. Apo) to mostly brown or dark brown (Bukidnon specimens). The ventral parts of all the examined specimens are extensively darkened. An examination of external morphological features and the male genitalia of specimens from these two areas confirmed that they belong to a single spe- cies. Angerianus palawanensis sp. n. (figs. 15-17) Type material. - Holotype @: Philippine Islands: Pa- 275 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Figs. 15-17. Male genitalia of Angerianus palawanensis. - 15, Vesica; 16, Left paramere, dorsal view; 17, Left paramere, postero- lateral view. lawan: Mantalingajan, Pinigisan, 600m, Sept. 6, 1961 (UZMC, Noona Dan Exp.). Diagnosis. - Recognized by the dark brown general coloration, with legs and antennae mostly pale; head with broad, longitudinal stripe between eyes; pronotum appearing glabrous, with scattered, fine punctures on collar and disc; length of labium much greater than posterior width of pronotum, and male genitalia as in figures 15-17. Description of male holotype. - Length 2.30; dark brown general coloration. Head: Width across eyes 0.66, vertex 0.31; pale brownish yellow, except dark brown posteriad of eyes ventrally and later- ally, and with fuscous stripe between eyes extend- ing to anterior margin of pronotum. Antennae: I, length 0.77, basal half dark yellowish brown, distal half slightly paler; II, length 1.59, pale brownish yellow; III & IV, brown. Labium: Length 1.31; reaching to posterior margin of metacoxae; seg- ments I & II dark brown, III & IV pale brownish yellow. Pronotum: Posterior width 1.06; collar and disc with scattered fine punctures, appearing gla- brous; uniformly dark brown, except somewhat paler laterally dorsad of coxal cleft; scutellum dark brown, less strongly elevated above level of hemel- ytra; scent gland ostiole dark brown, similar in coloration to surrounding pleura. Hemelytra: Cla- vus and corium dark brown, becoming slightly paler distally; cuneus brown basally, outer margin and apex broadly pale, almost clear; membrane lightly tinged with fuscous basally, veins brown, paler distally. Legs: Uniformly pale yellow. Geni- talia: Figures 15-17. Female. - Unknown. Etymology. - Named for its occurrence on the island of Palawan. Distribution. - Palawan, Philippine Islands. 276 Angerianus pallidus sp. n. (figs. 18-20) Type material. - Holotype @: Malaysia: Sarawak: Nanga Pelagus, nr. Kapit, 180-585 m, Aug. 7-14, 1958, T.C. Maa (BISH). — Paratype: Malaysia: Sabah: 1 &, Ranau, Sept. 30-Oct. 5, 1958, T.C. Maa (BISH). Diagnosis. - Similar to mindanao in size and general coloration, but distinguished by the broader head and vertex, weakly punctate pronotal disc, and structure of the male genitalia (figs. 18- 20). The pale general coloration and faint punc- tures on the pronotal disc will distinguish pallidus from fractus, maurus and palawanensis. Description of male. - Length 2.55-2.77; pale brownish yellow general coloration. Head: Width across eyes 0.73-0.77, vertex 0.36-0.39; uniformly brownish yellow. Antennae: I, length 0.88, pale brownish yellow; I-IV, missing. Labium: Length 1.39; reaching between mesocoxae; uniformly pale brownish yellow. Pronotum: Posterior width 1.38; brownish yellow; collar and disc with very faint, shallow punctures and short, pale setae; scutellum brown or pale grayish yellow, moderately elevated above level of resting hemelytra; scent gland osti- ole dirty yellow, only slightly paler than surround- ing pleura. Hemelytra: Clavus brown or grayish yellow; corium and cuneus clear pale yellow - co- rium of holotype lightly tinged with fuscous ba- sally, especially bordering claval suture; membrane clear, with slight yellow coloration bordering veins. Legs: Uniformly pale brownish yellow. Genitalia: Left paramere and vesica as in figures 18-20. Female. - Unknown. Etymology. - From the Latin, pallidus (pale), referring to the pale general coloration. Distribution. - Sabah and Sarawak, Malaysia. Discussion. - The paratype of this species is uniformly pale dorsally, while the holotype has the scutellum and clavus darker than the rest of the dorsum. All other features of the two specimens including the male genitalia are indistinguishable, although it is noted that the holotype is slightly larger than the paratype. ACKNOWLEDGMENTS I thank the following individuals and their re- spective institutions for providing specimens for study: Randall T. Schuh, American Museum of Natural History, New York (AMNH); Gordon M. Nishida, Bernice P. Bishop Museum, Honolulu (BISH); William R. Dolling, Natural History Mu- seum, London (NHML); Richard C. Froeschner and Thomas J. Henry, National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM); and Nils Meller Andersen, Univer- sitetets Zoologiske Museum, Copenhagen (UZMC). Bonnie B. Hall, Scientific Illustrator, Systematic Entomology Laboratory, Department of Entomol- ogy, Oregon State University prepared the dorsal habitus illustration of Angerianus maurus, with funds made available by the Oregon State Univer- sity Foundation (Systematic Entomology Labora- tory Fund). Assistance with preparation of the scanning electron micrographs was received from John Spratt and Susan Barnes, SEM Unit, NHML. REFERENCES Cassis, G., 1984. A systematic study of the subfamily Dicyphinae (Heteroptera: Miridae). - Ph. D. diss., Oregon State Univ., Corvallis, 389 p. STONEDAHL: Review of Angerianus Figs. 18-20. Male genitalia of Angerianus pallidus. — 18, Vesica; 19, Left paramere, dorsal view; 20, Left paramere, posterolateral view. Carvalho, J. C. M., 1952. On the major classification of the Miridae (Hemiptera). (With keys to subfamilies and tribes and a catalogue of the World genera). - Anais da Academia Brasileira de Cièncias 24: 31-110. Carvalho, J. C. M., 1955. Keys to the genera of Miridae of the World. - Boletim do Museu Paraense Emilio Goeldi 11: 1-151. Carvalho, J. C. M., 1957. Catalogue of the Miridae of the World. Part I. Subfamilies Cylapinae, Deraeocorinae and Bryocorinae. - Arquivos do Museu Nacional, Rio de Janeiro 44: 1-158. Distant, W. L., 1904. The Fauna of British India, Including Ceylon and Burma. Rhynchota. Vol. II (Heteroptera). - Taylor and Francis, London, 503 p. Henry, T. J., Neal Jr., J. W. & K. M. Gott, 1986. Stetho- conus japonicus (Heteroptera: Miridae): A predator of Stephanitis lace bugs newly discovered in the Uni- ted States, promising in the biocontrol of Azalea Lace Bug (Heteroptera: Tingidae). - Proceedings of the Entomological Society of Washington 88: 722-730. Kerzhner, I. M., 1967. Family Miridae (Capsidae). - In: G. Ya. Bei-Bienko (ed.), Keys to the Insects of the European USSR. Vol. I. Apterygota, Palaeoptera, He- mimetabola. - Zoological Institute, Academy of Scien- ces of the USSR, Leningrad. 913-1003. [English trans- lation by Israel Program for Scientific Translations, Jerusalem. Printed by S. Monson Binding, Jerusalem]. Reuter, O. M., 1910. Neue Beiträge zur Phylogenie und Systematik der Miriden nebst einleitenden Bemer- kungen tiber die Phylogenie der Heteropteren-Fami- lien. - Acta Societatis Scientiarum Fennicae 37(3): 1- 167. Stonedahl, G. M. & G. Cassis, 1991. Revision and cladistic analysis of the plant bug genus Fingulus Distant (He- teroptera: Miridae: Deraeocorinae). — Novitates, In Press. Received: 25 March 1991 Accepted: 2 July 1991 277 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Continued from p. 200 area. The ecological section deals with zonation of dung and carrion beetles and Lepidoptera, and in- cludes specialized studies on several groups of Co- leoptera, Hymenoptera, Diptera, Lepidoptera and aquatic Acari. A list of the publications related to the expedition concludes the volume. My opinion on this publication cannot be objec- tive. Working myself on the Odonata of Sulawesi, I am interested in every paper on this island. Of course I love to see the forest profiles of the places CORRIGENDA I have worked myself. The extensive contributions on the aquatic fauna make this publication even more attractive to me. On the other hand, this book presents so many new and interesting data, that I am convinced that it deserves a broader readership than the expedition members alone. I can warmly recommend it to anybody interested in tropical entomology. The book is good value for money, although the quality of typesetting and printing is below one would expect for the final presentation of such an important enterprise. [J. van Tol] Gielis, C., 1991. Pterophorus Schäffer in the Ethiopian region (Lepidoptera: Pteropho- ridae: Pterophorinae): new species and checklist. - Tijdschrift voor Entomologie 134: 1- 8, figs. 1-21. The editors regret that the plates on pages 4 and 6, in the above mentioned paper, showing the fe- male genitalia, were incorrectly composed, so that the figure numbers for figs. 10-20 were wrongly assigned. The following key gives the correct figure numbers: 10 change into: 15 El change into: 16 12 change into: 17 13 change into: 18 14 changeinto: 19 15 change into: 20 16 change into: 10 17 changeinto: 11 18 changeinto: 12 19 changeinto: 15 20 change into: 14 21 remains: 21 For convenience, we also include adapted cap- tions for page 4 and 6 respectively: page 4: Figs. 15-20. - Pterophorus spp., female genitalia. - 15, P. candidalis (Walker), Côte d'Ivoire, slide CG 2182 (CG); 16, P. candidalis (Walker), Chad, slide CG 2186 (cG); 17, Idem, Tanzania, slide cc 4020 (zMuc); 18, P. dallastai sp. n., paratype, slide CG 1851 (MRAC); 19, P. massai sp. n., holotype, 20; P. wzungwe sp. n., paratype, slide CG 4040 (ZMUC); page 6: Figs. 10-14. - 21. Pterophorus spp., female genitalia. 10, P. alibidus (Zeller), Tanzania, slide CG 4018 (zMuc); 11, P. rhyparias (Meyrick), Zaire, slide Bigot (MRAC); 12, P. cerawnia (Bigot), Benin, slide CG 1881 (MNHN); 13, P. lampra (Bigot), Côte d'Ivoire, slide cG 2178 (CG); 14, P. spissa (Bigot), paratype, Zaire, slide Bigot (MRAC); 21, P. cleronoma Meyrick, paralectotype, slide Bigot (MNHN). 278 H. TURIN, K. ALDERS, P. J. DEN BOER, S. VAN ESSEN, TH. HEIJERMAN, W. LAANE & E. PENTERMAN Dutch Carabidological Association ECOLOGICAL CHARACTERIZATION OF CARABID SPECIES (COLEOPTERA, CARABIDAE) IN THE NETHERLANDS FROM THIRTY YEARS OF PITFALL SAMPLING Turin, H., K. Alders, P. J. den Boer, S. van Essen, Th. Heijerman, W. Laane & E. Penterman, 1991. Ecological characterization of carabid species (Coleoptera, Carabidae) in the Netherlands from thirty years of pitfall sampling. - Tijdschrift voor Entomologie 134: 279-304, figs. 1-20, tabs. 1-10. [ISSN 0040-7496]. Published 18 December 1991. An ecological characterization of 285 Dutch carabid species based on a large data set obtained by pitfall trapping in various habitats all over The Netherlands is presented. The data set contains ca. 1.5 million specimens, collected during thirty years in 1616 year- samples from 862 localities. Information about habitat, viz. type of vegetation, type of soil and humidity was recorded for each of the 862 sampling localities. Two Way Indicator Species Analysis (TWINSPAN) as well as Detrended Correspondence Analysis (DECORANA) were used to classify the species into habitat groups. Seven principal habitat groups could be recognized. Indices for ecological amplitude, preference for soil type and humidity, are formulated and estimated for each species. This resulted in a description of the habitat preferences of the Dutch carabid species in far greater detail than was possible before. The classification is compared with others from the literature. Correspondence: H. Turin, Esdoorndreef 29, 6871 LK Renkum. The Netherlands. Key words. - Ecology, Carabidae, Twinspan, Decorana, pitfall In 1945, Lindroth published the first volume of his major work ‘Die Fennoskandischen Carabidae’ (Lindroth 1945). In this work he described the ecological preferences of the Fennoscandian ca- rabid species. Since pitfall trapping was not prac- ticed until about 1950, Lindroth’s ecological char- acterizations were based on his own hand- collections, supplemented by those of other collec- tors, on laboratory experiments and on data taken from literature. This resulted in detailed descrip- tions of the species’ ecological preferences. In 1949, Lindroth classified the species into a number of ecological groups, recognizing xerophilous species, mesophilous species (including ubiquists), hygro- philous species, arboreal species, forest species and synanthropous species (Lindroth 1949). From about 1950 onwards, pitfall trapping be- came a commonly used technique in Europe. Thiele (1977) gave a summary of many ecological studies based on pitfall trapping carried out until ca. 1975. On basis of the results of these studies he charac- terized the carabid faunas of certain types of hab- itat, such as forests, sandy habitats and cultivated land, by presenting lists of characteristic species of these habitat categories. However, his tables only show the most abundant species in that particular habitat group giving no information about the oc- currences of these ‘typical’ species in other types of habitat. Luff et al. (1989) were the first to classify hab- itats of ground beetles based on a large data set from nearly 250 sites in North-East England. The carabid samples were grouped, based on pres- ence/absence data, using the Two Way Indicator Species Analysis (TWINSPAN (Hill 1979a) and ordi- nated by Detrended Correspondence Analysis (DECORANA (Hill 1979b). Ten primary groups of carabid habitats were recognized, viz.: coastal, up- land (dry, wet), woodland, grassland (dry, wet), riverside (boulders, shingle, sand) and marsh. Eyre & Luff (1990a) made a preliminary classification of European grassland habitats using carabids. The carabid assemblages of 363 pitfalled grasslands throughout Britain, were described by Eyre & Luff (1990b) The characterizations of Lindroth (1945, 1949) 279) TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 are usually considered to be accurate, although they were obtained with a non-quantitative method. Furthermore, Lindroths classification applies to the Fennoscandian situation, and species preferen- ces may show geographical variation. The major disadvantage of the analyses of Thiele (1977) is, that his compilations are based on data from the British Isles to Russia. Many of the spe- cies only occur in a limited part of this area. The present study was carried out because there is a need for adequate ecological characterizations for the Dutch area with respect to future work, especially in the field of nature management and conservation. The material that is available from The Netherlands allows us to use quantitative me- thods to describe the ecological preferences of 285 species in our area in great detail. The ecological characterizations can serve as a basis for future descriptions and evaluations of carabid faunas of specific areas within The Netherlands. MATERIAL AND METHODS Material When preparing the first edition of the Dutch carabid atlas (Turin, et al. 1977) it appeared that an enormous amount of data from pitfall trapping was available. Pitfalls were used in The Nether- lands already in the early 1950’s by Van der Drift, soon followed by Den Boer in 1953. In the past, several studies concerning the effi- ciency of pitfall sampling have been carried out. A summary of the results of these investigations up to 1975 was published by Thiele (1977), concluding that pitfall trapping is a suitable technique for investigating carabid populations in a quantitative way. Den Boer (1977, 1990) reviewed investiga- tions carried out by pitfall sampling at the Biolog- ical Station Wijster. It appeared that so-called year- catches of most carabid species give reliable rela- tive estimates of the mean densities of active adult specimens around the pitfalls (Baars 1979, Den Boer 1979). A year-catch is defined as the summa- tion per species of all specimens caught in one series of pitfalls during one year (or at least during the main reproductive period of carabids). These estimates are relative, meaning that they can be used only to compare the relative abundances within a certain species over a series of samples and/or years, but not between species. This is be- cause each species has characteristic ‘catch-parame- ters’, such as activity pattern, way of living and catchability (see Luff 1975, Den Boer 1986). Fig 1. The number of sampled sites per 10 km square of the UTM-grid in the Netherlands. 280 The dataset for the present study consists of 1616 year-samples from 862 localities in The Ne- therlands (fig. 1), covering the period 1953-1983. A list of all year-samples included in the database is given by Turin & Penterman (1985). The number of pitfalls used in a pitfall series and the sizes of the traps varied considerably between investigators. Therefore, the number of specimens in a year-sample were standardized by calculating the number of specimens per decimeter pitfall edge per year (SDY). Before considering the catches, all sampling lo- calities were uniformly described, using the ‘eco- code’ of the European Invertebrate Survey for The Netherlands (Van Tol 1979) ina slightly modified version (Penterman & Turin 1985). In this code information about type of vegetation, soil type, soil humidity, size of the locality and type of manage- ment is recorded. This information allowed us to recognize 33 habitats (table 1, p. 292), according to which the 1616 year-samples were coded. MEAN 1n(SDY+1) Fig. 2. The frequency distribution of the species mean In(SDY+1)-values over each of the 33 habitats, taken from the data matrix. A: the Y-axis shows the number of relative abundances (see text) in the data matrix. B: the mean In(SDY+1)-values; only a very small part of the X- axis is shown. This means that the values are very low all over the data matrix. These figures proved to be unwor- kable for TWINSPAN classification. A. 1600 PERCENTUAL VALUES Fig. 3. The distribution of the pertage values after rescal- ing the mean abundances per species, with the maximum mean In(SDY+1) put to 100% (relative occurrences). A: the Y-axis shows the number of relative occurrences. All figures from all 33 habitats are included except the first highest percentage value (* = 100%) of each species, which was used as a basic value for the rescaling. B: the percentage values, where: .=<5%, 1 =5-15%, 2 = 15- 25% etc., 9 = 85-95% and * = > 95%; for further explanation see text and tables 2-9. TURIN ET AL.: Carabid beetles Classification of habitats and habitat preferences of species Catches of a single carabid species from a large number of pitfall samples over many years, tend to form a log-normal distribution (Den Boer 1977). Therefore, the number of specimens per decimeter per year (SDY), was transformed to natural loga- rithms. For each of the 33 habitat categories (table 1) the mean value of In(SDY+1) was calculated per species. Since all year-samples contribute to the mean In(SDY+1) of a species, including those where the species was not recorded or in very low numbers only, the resulting values were very low for the majority of the species (fig. 2). Therefore, the values for the relative abundances were re- scaled, by setting the highest value for a species at 100% and recalculating the SDY-values for the other habitats accordingly (fig. 3), in this way transforming the relative abundances into relative occurrences. This also made the catches of different species better comparable. We did not rescale the abundances of the species per habitat, in order to save the differences in significance between habi- tats as suitable sites for survival. TWINSPAN was applied to the relative occurren- ces of the species, to classify the habitats into hab- itat groups and to obtain a classification of the species according to their habitat preferences based on the rescaled mean In(SDY+1)-values. TWINSPAN was used using cut-levels 20, 40, 60 and 80% and was run several times using different pseudospecies’ cut-levels. The results of these runs were all very similar when using the rescaled mean In(SDY+1)-values. The results with different cut- levels based on the mean In(SDY+1) values with- out rescaling, however, differed considerably among each other and were not further used. The resulting classification of habitats will be presented in the form of hierarchical tree diagrams and scat- tergrams. The full two-way table is presented, showing the group division of the species and con- taining the values indicating the relative occurren- ces as defined by the pseudospecies’ cut levels. The information presented in this table allows us to easily evaluate the habitat specificity of a particular species. Ecological amplitude Two indices were defined and calculated that estimate the ecological amplitude of a species. The first measure (PRES) is based on the species pres- ences over habitats, and is defined as: n. PRES, = —1— n 281 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 where PRES; represents the ecological amplitude of the j-th species, n; the number of habitats in which this species is present, and n the total number of habitats recognized (33). PRES will range from 0 to 1. For the second ‘eurytopy’ estimate (SIM) the index of Simpson (D) is used (Simpson 1949), which characterizes the distribution of the abun- dances over the habitats in the form of: 1-D. This may thus be written as: where a, is a Measure of the relative abundance of a species in habitat group j and is equal to mean In(SDY+1),, while n CE aT > In (SDY + 1); aa The value of SIM may range from 0 to 1. Soil type and humidity When characterizing the sampling localities, three humidity classes were recognized, viz. ‘dry’, ‘moist’ and ‘wet’. The humidity preference of a species was estimated at an ordinal scale from 1 (very xerophilous) to 5 (very hygrophilous). Spe- cies only caught in ‘wet’ localities score 5, species from ‘wet’ as well as from ‘moist’ localities score 4, species only from ‘moist’ localities score 3, species from ‘moist’ as well as from ‘dry’ localities score 2 and species only from ‘dry’ localities score 1. Each sampling locality was also classified into one of seven soil types: limestone (li), loam (lo), loamy sand/sandy clay (ls), river clay (rc), sea clay (sc), peat moor (pm) and sand (s). A species is regarded to show a preference for a certain kind of soil when the In(SDY+1), averaged over all local- ities with this specific soil type, is at least 2 times the sum of the mean In(SDY+1) values of the localities from the other soil types taken together. Nomenclature and taxonomy The nomenclature follows Turin (1990), except for three Calathus species of the melanocephalus group, for which is referred to Aukema (1990). The material from pitfall trapping presented here un- der the name of C. melanocephalus in fact is a mixture of C. melanocephalus and C. cinctus (Au- kema 1990). Only in the more recent sampling (ca. after 1980), C. cinctus was recognized as a separate 282 species. A similar problem exists where it concerns the species Pterostichus nigrita and Asaphidion flavipes. Pterostichus rhaeticus was not separated from P. nigrita (see: Koch 1984), and Asaphidion curtum as well as A. stierlini not from A. flavipes (see: Lohse 1983, Lompe 1989, Schweiger 1975). The full names of all species are given in the ap- pendix. RESULTS Classification of habitats Figs. 4-11 present the results from the TWINSPAN classification of the relative occurrences over the 33 habitats from table 1. Seven main habitat groups could be recognized (fig. 4): Group I. - Peat and heath vegetations, habitats 1-5. A further subdivision of this end-group is presented in fig. 5. Group II. - Poor grassland and dune habitats, habitats 6-11 (subdivision see fig. 6). Group III. - Cultivated land and open coniferous plantations, habitats 12-15 (subdivision see fig. 7). Group IV. - Mature forests, habitats 16-20 (sub- division see fig. 8). Group V. - Moist/wet forests (forests in water meadows; brook forests) and ruderal grass locali- ties, including limestone grassland, habitats 21-25 (subdivision see fig. 9). Group VI. - Moist habitats overgrown with weeds, polder-(colonization-) habitats, habitats 26- 30 (subdivision see fig. 10). Group VII. - Wet habitats/shores, habitat 31-33 (subdivision see fig. 11). Fig. 4. Dendrogram of the main division into seven groups according to TWINSPAN classification of the spe- cies’ relative occurrences. I-VII = main habitat groups: I = Peat and heathland vegetations (habitat 1-5, see fig. 5), II = Poor grassland and dune habitats (habitat 6-11, see fig. 6), III = Cultivated land and open coniferous planta- tions (habitat 12-15, see fig. 7), IV = Mature forests (habitat 16-20, see fig. 8), V = Moist / wet forests and ruderal grass localities, including limestone grassland (habitat 21-25, see fig. 9), VI= Moist habitats, overgrown with weeds, polder-(colonization-) sites (habitat 26-30, see fig. 10), VII = Wet habitats/shores (habitat 31-33, see fig. 11). 1 PEAT MOOR MOLINIA HEATH, ERICA 4 HEATH, CALLUNA 5 HEATH, DESCHAMPSIA m 5 | 6. CORYNEPHORETUM 7 COASTAL DUNES 8 DUNE GRASSLAND [soi DUNE FOREST 10 DUNE SCRUB 11 POOR GRASSLAND 12 GRASSLAND, PASTURES 13 ARABLE LAND 14 WASTE LAND 15 CONIFER. PLANTATION 16 CONIFEROUS, MATURE 17 CONIFEROUS, MOIST OAK BIRCH OAK BEECH OAK HORNBEAM 21 POPLAR, MOIST 22 ALDER WILLOW INLAND SCRUB, MOIST RUDERAL, PARKS ETC. | 25] LIMESTONE GRASSLAND | 26 | GRASSLAND, HERBS 27 REEDLAND, SAND SILT 28 REEDLAND, POLDERS 29 ARABLE, POLDERS 30 COLONIZATION SITES 10 SEASHORE, SAND BARS 32 RIVER BANKS, OPEN 33 SALT MARSHES 11 Figs. 5-11. Subdivision of habitat groups I-VII (see fig. 4 and table 1). - 5, group I; 6, group II; 7, group III; 8, group IV; 9, group V; 10, group VI; 11, group VII. TURIN ET AL.: Carabid beetles Fig. 12 shows the results of an ordination of the relative occurrences over the habitats by DECORANA, where the habitat groups mentioned above are indicated with dashed lines. The TWINSPAN divisions form rather coherent groups. The first axis of DECORANA is clearly correlated with moisture, dry habitats: 15-16, coniferous forest; 6, Corynephoretum; 4, Calluna-heath on the left, and wet: 31-33, shore habitats on the right. The second axis seems to be related in some way with the structure of the vegetation, open vegeta- tions: 1-5, heath and peat habitats, 6-11, dune hab- itats in the lower part, and highly shaded vegeta- tions: 16-22, forests in the upper part of the figure. Classification of species The habitat preferences of the species are indi- cated by their relative occurrences over the 33 hab- itats. The species are divided into eight species groups: A-H (fig. 13), and are tabulated in tables 2-9. Most groups contained very eurytopic as well as rare species. These species have been taken from the original groups and are brought together into two separate tables: 8 (eurytopic species) and 9 (rare species). Species group A (table 2): Species of heath vege- tations and peat moor, mainly occurring in habitat group I. Species group B (table 3): Species of sandy local- ities, such as dunes, arable land and coniferous plantations; principal occurrences in habitat-group II and/or III. 500 200 n Nees o! de” degl ___ Zen. à 100 200 300 400 500 Fig. 12. Ordination by DECORANA, presenting on the first two axes the projections of the species scores for the 33 habitats (explanation see table 1). The dashed lines indi- cate the main habitat-groups: 1-5 = peat and heathland vegetations, 6-11 = poor grasslands and dune habitats, 12-15 = cultivated land and open coniferous plantations, 16-20 = mature forests, 21-25 = moist to wet forests and ruderal grassland habitats, including limestone grassland, 26-30 = moist habitats, reedland, pioneer (colonization- ) habitats, polders, 31-33 = wet habitats, shores and river banks. 283 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 1000, 1001, 1010, 1011 1110, 1111 Es Fig. 13. Dendrogram of the species groups according to the TWINSPAN classification. The letters A-H indicate the TWINSPAN end-groups, treated in the respective species group. The numbers 0000-1111 indicate the first four levels of the TWINSPAN classification (compare table 2-7). 1100, 1101, Species group C (table 4): Species of open areas, dune localities as well as colonization sites and shores, mainly in habitat-groups II, III and/or VI, VII. Species group D (table 5): Species of forests and/or ruderal places, including limestone grass- land; occurrences mainly in IV and/or V. Species group E, F, G (table 6): These groups are not treated separately, because the separate end- groups are rather heterogeneous and less typical for a certain habitat group than the other species- groups. However, many species occur in moist and shaded localities, such as moist forests, scrubs and reedland; having an accent in habitat groups V and VI, and others show no special preference for moist habitats. Species group H (table 7): Species of more open moist-wet habitats, reedland and shores; occurren- ces mainly in groups VI and/or VII, (TWINSPAN species end-groups 1100, 1101, 1110 and 1111). Eurytopic species (EU) (table 8): These species are placed in a separate group on basis of their ecological amplitude: PRES > 0.75 or SIM > 0.85. The species are arranged according to their eury- topy estimates (Pres, Sim). The end-group number indicates from which of the species groups menti- oned above (A-H) they originate. Rare species (R) (table 9): Species have been placed in this group when the number of samples (Sa) < 6 and when the number of individuals < 50. If the species is merely present in 3-5 year-samples and these samples belong to the same habitat, the species was not placed into group H. In tables 2-9 relative occurrences are presented 284 according to the percentual scaling, where for each species the highest mean In(SDY+1) in one of the 33 habitats, is put at 100%. In the body of the tables the numbers indicate: 1 = 5-15%;2= 15-25% etc., 9 = 85-95% and * = > 95% respectively. Points indicate the habitats where the species is present, but where the relative occurrences are below 5%. Species were ordinated by DECORANA, according to their relative occurrences per habitat. In figs. 14- 20 the position of the species belonging to the groups A-H on the first two axes of the ordination is given. The first axis shows a dry-wet gradient: species preferring dry habitats (group A) on the left (fig. 14), and species from wet habitats (group H) on the right (fig. 19). The second axis is related with amount of shade: species of exposed habitats (group A) at the bottom (figs. 14, 15), and forest dwelling species (group D) at the top of the figure (fig. 17). The highly eurytopic species of group EU, take a central position according to that of the other species groups, resembling that of the inter- mediary groups E, F and G (fig. 18). Ecological amplitude, soil type and humidity The indices for the ecological amplitude of the species (Pres, Sim), for soil preference (So) and humidity preference (Hu) are given in the right columns of the tables 2-9. Also the number of year- samples (Sa), the number of individuals (N). DISCUSSION OF THE SPECIES GROUPS In this chapter we will give a short discussion per species group. Analyses more into detail can be expected in further papers where the species com- position of several habitats or habitat-groups will be treated. Species-group A (table 2 p. 294) Most species listed in table 2 show clear prefer- ences for peaty soils and/or sandy soils. When they have no preference for any soil type, they are mainly restricted to oligotrophic habitats. The hu- midity preference varies: most species that also occur in the Corynephoretum, such as Pterostichus lepidus, Bradycellus ruficollis and Amara infima, do not occur in humid habitats. Other species have a rather high preference for humid sites (humidity 4- 5), such as Pterostichus aterrimus, Agonum ericeti, and Anisodactylus nemorivagus, which are known from literature as true peat-moor dwellers (Lin- droth 1945, Mossakowski 1970a, 1970b). The re- sults agree with those of investigations in German heath-moor complexes (Horion & Hoch 1954, Grossecappenberg et al. 1978). According to the TWINSPAN-classification (figs. 4-5), peat moor has Fig. 14-20. Ordination of the species groups A-H and group EU, by DECORANA. - 14, group A: species of heath vegetations and peat moor (see table 2); 15, group B: species of sandy localities, dunes, arable land and conif- erous plantations (see table 3); 16, group C: species of open areas (see table 4); 17, group D: forest species and species of ruderal places including limestone grasslands (see table 5); 18, groups E, F and G: species occurring in most and shaded habitats (see table 6); 19, group H: species of moist-wet habitats, reedland and shores (see table 7); 20, group EU: eurytopic species (see table 8); the outlines of the previous groups are indicated to accentu- ate the central position of the eurytopic species. been included in the present species group and not in one of the moist groups E or F, in spite of the occurrence of many moist-preferring species. This can be explained by the fact that the fauna of peat- moor is rather poor in species and that the majority of these species is ecologically more related to heath-like vegetations. The peat-moor fauna has hardly any species in common with the carabid assemblages of shore habitats (species-group F), where the occurrence of many Bembidion-species is most characteristic. The results in table 2 agree well with those from the literature. Some studies carried out in Germany by Mossakowski (1970a, 1970b) and Rabeler (1947) also recognize Agonum ericeti, Anisodactylus nemorivagus, Bradycellus ru- TURIN ET AL.: Carabid beetles ficollis and Trichocellus cognatus as characteristic species for peaty soil. Mossakowski (1964, 1970c) and Rabeler (1947) mention some species charac- teristic for heath vegetations, such as: Amara in- fima, B. ruficollis, Carabus arvensis, C. nitens, Cic- indela campestris, Olisthopus rotundatus and Pterostichus lepidus. Some of the species menti- oned by these authors can be found in the ecolog- ically most related species group B (table 3) or they are arranged among the eurytopic or rare species (tables 8-9), e.g. Bembidion nigricorne, Bradycellus caucasicus, Cicindela sylvatica, Pterostichus versic- olor, Syntomus foveatus as well as several Calathus and Cymindis species. Species group B (table 3, p. 294 ) This group is divided into two subgroups, based on a sixth level TWINSPAN division: Bl - species mainly occurring in the Corynephoretum, dune habitats and poor grassland; B2 - species of sandy arable land, waste land and young coniferous plan- tations on sand. The species of group Bl are in general confined to poor, dry and open, sandy hab- itats especially in dune areas. Some species are very stenotopic and more or less restricted to drifting sands of coastal and inland dunes: Harpalus servus, Harpalus vernalis, Masoreus wetterhali, Ophonus cordatus and Harpalus neglectus. Some papers con- cerning dunes and Corynephoretum sites (Schjatz- Christensen 1957, 1966a, 1966b) mention several Harpalus species e.g.: H. anxius, H. solitaris, H. neglectus and H. smaragdinus as being typical for dry and sandy areas in Denmark. The following species can also be found in heath vegetations and they are obviously ecologically related to the spe- cies of group A: Bembidion nigricorne, Cymindis macularis, Notiophilus germinyi and Cicindela syl- vatica. Most species in the Bl group have low humidity indices (1-2), and are rather stenotopic. The species belonging to group B2 are character- istic for many types of sandy habitats. As far as they occur in cultivated areas, most of them avoid heavy fertilization. Species common to intensively culti- vated agricultural land, can be found among the eurytopic species (see table 8). Laemostenes terric- ola, Calathus micropterus and Pterostichus quadri- foveolatus are also related to forests (habitat group IV). Species group C (table 4, p. 295) This small group consists of four species that occur primarily in open localities, dry as well as rather wet ones. Clivina collaris is predominantly a riparian species. The other species are confined to open country with a sparse vegetation and show a preference for coastal areas; this especially app- lies to Calathus mollis. 285 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Species group D (table 5, p. 296) Three subgroups were grouped together based on their occurrences in the main habitat groups IV and V: DI - Stenotopic species of cultivated and ruderal sites such as poor unfertilized fields, gardens and limestone grasslands, mainly on li- mestone soil in the southern part of the province of Limburg; D2 - Species of more or less moist and shaded places partly on limestone soil; D3 - Eury- topic and stenotopic forest species. Belonging to the first subgroup are several heath-preferring species (xerotherm species: Becker 1975, Lindroth 1949) which in The Nether- lands are exclusively confined to limestone soil: Parophonus maculicornis, Amara nitida, Brachinus crepitans, Harpalus dimidiatus, and Ophonus mel- leti. Other thermophilous species can also be found on localities outside the limestone area of The Netherlands, on open sun-exposed, xerotherm sites such as slopes of river dikes with a southern exposition (Turin 1983, Turin et al. 1977): Amara montivaga, Ophonus puncticeps, Harpalus ru- bripes, and Lebia chlorocephala. Ophonus ruftbar- bis is the least thermophilous and most eurytopic Ophonus species. It can be found on more or less shady sites (Lindroth 1974, 1986). The species of the D2 subgroup are restricted to the southern part of the province of Limburg or have a more or less fluviatile distribution in The Netherlands. Most species prefer chalky, clayish soil or loam, some of them inhabiting shaded (wooded) sites: Stomis pumicatus, Bradycellus sharpt, Carabus coriaceus, and Pterostichus madi- dus. These species also occur in limestone grass- land at slopes with a northern exposition (Turin 1983). Subgroup D3 consists of the true forest species. The first six species are rather eurytopic, which also applies to Lezstus rufomarginatus and Carabus problematicus. Some of the more stenotopic forest species are more or less restricted to the oak-horn- beam forests of the southern part of Limburg and the easternmost part of The Netherlands, e.g. Abax parallelus, Molops piceus, and Trichotichnus nitens. Amara praetermissa can be found in the Corynephoretum, but also at acid sites on top of the limestone hills in Limburg with a more heather- like (Calluna) vegetation, which explains the pres- ence of this species in the D-group. There are far more investigations into the com- position of the carabid fauna of forests than in that of heathland vegetations. Nowadays heathland is rapidly disappearing from The Netherlands, but about a hundred years ago it was one of the most prominent components of the landscape. In 1850 the area covered with forest was below 3%. Affo- restation, which started in the first part of this 286 century, has increased this area to about 6% in 1950 and 8-9% at the moment. Thus, many Dutch forests are rather young. The majority of this fore- sted area consists of coniferous and exotic trees. Apart from the light oak-birch forests, which form the more natural climax vegetation on sandy soils, and the riverine forests, the more rich types of deciduous forest, e.g. Querco-Carpinetum, can only be found in restricted areas. Species characteristic (Thiele 1977) of these kind of forests in Central Europe, e.g. Abax ovalis (not in pitfalls), Carabus auronitens, Molops piceus, Pterostichus cristatus and Trichotichnus nitens, are restricted to the very eastern and southern parts of The Netherlands. Thiele (1977) reviews studies on the most abund- ant species of the (sub)centreuropic forests, includ- ing many investigations from Germany, such as those of Heckendorf et al. (1986), Kolbe (1968, 1970), Lauterbach (1964), Rabeler (1957, 1962, 1963, 1967 and 1969) and Thiele (1956). Also li- terature concerning The Netherlands was taken into account by Thiele, e.g. Van der Drift (1959) and Den Boer (1965). Recently, Turin & Heijer- man (1988) carried out a first survey on the present data in a more simple way by only examining the species having the highest numbers of individuals and presences in all types of forest in The Nether- lands together, and arranging them according to the fraction of specimens caught in forests, as com- pared with those in other habitats. The terms ‘stenotopic / eurytopic forest species’ they use, only apply to the Dutch area. Especially the species of mountainous localities are missing (highest alti- tude in The Netherlands is below 400 m). Most stenotopic forest species mentioned by Turin & Heijerman are now listed in table 5. Heijerman & Turin (1989) found great differences in fauna com- position between forests in different parts of The Netherlands. This can possibly be explained by the fact that most of the forested areas are rather young and have been colonized recently. Species group E, F, G (table 6, p. 297) This group is polythetic and in fact a compilation of seven small TWINSPAN end-groups. The species predominantly occur in moist and shaded habitats. Most species are hygrophilous, reaching high numbers in wet grassy forests, moist grasslands and reedland: El - species of moist forests, some- times in dune valleys and in moist grassland; F1 - two species of rather open and dynamic habitats; F2 - species of moist and shaded sites; Gl - only a single species; G2 - species with high numbers in reedland and in moist grassland, Oodes helo- pioides lives at wet and shaded sites; G3 - only Bembidion obtusum, which more or less prefers cultivated country; G4 - mainly hygrophilous spe- cies of wet meadows, marshes and reedland. Most species in group G4 inhabit rather wet and dense, shaded vegetations. Exceptions are: Amara ovata and A. famelica, which both live in more open sites. Investigations into the fauna of moist and wet habitats are for reedland, swamps and lake shores: David & Marchal (1963), Dawson (1965), Jarmer (1971), Koch (1977), Obrtel (1972), Renkonen (1944) and Wasner (1977). The present results largely agree with the studies mentioned. Species group H (table 7, p. 298) The species belonging to group H reach high numbers in the moist or wet habitat groups VI and VII. Subdivisions based on the TWINSPAN classifi- cation are: Hl - a single rather eurytopic, coloniz- ing species: Amara similata, H2 — mostly hygro- philous species of young habitats in polders and in other colonization sites; H3 - four species of open sites near water; H4 - Species of shore habitats, a number of them confined to saline localities. Pter- ostichus cupreus is similar to Amara similata in living in moist grassland and arable land, both species being rather eurytopic. Bembidion bruxel- lense, Chlaenius nigricornis, Agonum albipes, Elaphrus cupreus and E. riparius are characteristic of river meadows and reedland. Amara convextus- cula is a typical species of sea clay soil and found mostly along the coast, but it occurs, just as Acupal- pus exiguus, Lastotrechus discus and Amara majus- cula, also in high numbers in the Dutch IJsselmeer polders (Turin et al. 1977). For large areas such as the Dutch polders, only constantly macropterous species and full-winged specimens of wing-dimor- phic species are capable of colonizing such areas (Haeck 1971, Meijer 1973, 1974, 1980). In colon- ization of small sites, such as burnings and clear- ings, also big wingless species may be able to im- migrate (Szyszko 1986), but in general, pioneer communities can be characterized by their high numbers of winged species. An example of long- term colonization and succession of carabids along rather homogenous road verges on clay soil in three Dutch polders of different age is given by Haeck et al. (1980). It illustrates that the carabid fauna of the Dutch polders keeps a coastal charac- ter for many decades. Most species of the H4-group live close to water in open habitats. Several species of this group are restricted to the salt marshes of the Waddensea area and the islands of the province of Zeeland: Dichetrotrichus gustavi, D. obsoletus, Pogonus chalceus, Bembidion normannum, B. pallidipenne, Dyschirius salinus, D. obscurus, and Bradycellus distinctus. About saline habitats several studies are available: Heydemann (1962, 1967), Mossakowski (1971), Niemela (1988) and Rueda & Montes TURIN ET AL.: Carabid beetles (1987) and they mention several species as typical for this habitat. The species occurring in saline habitats in The Netherlands are all treated in Turin (1991). Many of the characteristic and abundant species of inland shores and river banks cannot be found in Group H, since pitfalls do not function well in shore habitats. Especially focused on the riparian fauna of running water are the studies of: Andersen (1970, 1982, 1983), Krogerus (1948), Lehmann (1965) and Plachter (1986). Species living so close to the water that they hardly can be trapped with pitfalls are for instance: Agonum micans, Bembid- ion doris, B. obliquum, Dyschirius aeneus and Ne- bria livida. The species Bembidion articulatum, B. punctulatum, B. semipunctatum and B. testaceum, all from river banks and often abundant in The Netherlands, are not present in the pitfall material. Some shore species that also can be found at some distance of the water, are listed among the rare species (table 9). Eurytopic species (table 8, p. 299) A group of 53 eurytopic species and ubiquists, extracted from all habitat groups on basis of their ecological amplitude. Most species in this group show high occurrences and abundances in one of the habitat groups III, V and/or VI. Apart from a few species, they do not clearly prefer a special soil type or humidity class. Some of them have high tolerances with respect to fertilizers and intensive agricultural management and can therefore be nu- merous in arable land and in pastures. Thiele (1977), who compared the carabid faunas of arable land, meadows / pastures and clover / alfalfa, in fact gives incomplete information of the species preferences. The species listed in Thiele’s compi- lation are not characteristic for cultivated areas, but most of them simply are highly eurytopic species (see table 8). A similar survey was published by Basedow et al. (1976) and the results resemble those of Thiele. Notiophilus biguttatus and Nebria brevicollis are eurytopic forest species that also occur in dense, shady grasslands and in other shaded sites, such as gardens and orchards (Turin & Heijerman 1988). Rare species (table 9, p. 300-301) A groupof 59 rare species. Just as in the previous groups, the original TWINSPAN end-groups (GR) are mentioned after the species names. Twenty-six of the species only occur in one kind of habitat. For some species a lack of data is the main cause that the information on the species preferences is in- complete. However, for many species that have a highly restricted distribution in The Netherlands, especially those living near the fringes of their 287 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 ranges, the relative occurrences shown in table 9 will give a reliable picture of the species preferen- ces in the Dutch territory. The species of this group will be treated in a separate study. DISCUSSION Only a few studies concerning classification of carabid species based on a large dataset are known from western and north-western Europe. The most important are chronologically: Lindroth (1945, 1949), Thiele (1977), Den Boer (1977), Luff et al. (1989) and Eyre & Luff (1990a, 1990b). Lindroth (1945, 1949) indicated species prefer- ences for humidity, and certain vegetation types, such as woodland. Although his conclusions were based mainly on material from Fennoscandia, his classification proved to be valuable for a much larger area. It was used in many carabidological studies all over northern and western Europe. When comparing the results of the present study with Lindroth’s classification (table 10), one can conclude that they correlate rather well. The xero- philous species (X) in groups Al, Bl, B2, C1 and D1 (table 10), the hygrophilous species (H) in groups A1, G4, H2, H3 and H4 (table 10) and the forest species (HW+W-+WA) of Lindroth are found in groups D2, D3 and El. Lindroth’s meso- philous and more or less ubiquitous species (N) are scattered over a wide range of our species-groups with the highest numbers in the eurytopic EU- group (table 10). Fifty-nine species mentioned by Lindroth were caught in pitfalls in too low numbers for a characterization (group R) and 85 of the Dutch species mentioned by Lindroth, were not caught in pitfalls at all in The Netherlands (table 10 group O). From Lindroth’s characterizations (table 10) we can conclude that these species that could not be characterized in the present study, belong to the inhabitants of shores and river banks (80 species, table 10). Others are very rare or even extinct in The Netherlands, e.g. many xerophilous species (see Desender & Turin 1989, Turin 1990). Thiele (1977) attempted to characterize the carabid faunas of certain habitat groups such as forests, cultivated land and sandy habitats, compar- ing them on basis of species presence. He mainly used literature sources from the entire northwest- ern part of Europe. Although he included several lowland and mountain forests for the characteriza- tion of the forest fauna, we feel that this method gives an incorrect picture of the species preferen- ces, especially because the relative occurrences and abundances in other localities were not taken into account. Den Boer (1977) gives a far more complete pic- ture for 75 of the most abundant species in his 288 study area, by listing the data of hundreds of year- catches in many kinds of habitat in the province of Drenthe (The Netherlands). However, his study area is rather limited and many Dutch habitats and soil types are absent from his study (e. g. coastal dunes, clay soil and limestone grasslands). A com- parison with Den Boer’s results is not appropriate here, because his data form a significant part of our data set. Several extensive studies on classification of ca- rabid species have been published recently. Luff et al. (1989) is related to the classification of carabid habitats in north-east England, while another deals with the ground beetles of grassland habitats in Europe (Eyre & Luff 1990a). Finally, Eyre & Luff (1990b) presented a classification of ground beetles of the British grasslands in more detail. In all stu- dies DECORANA and TWINSPAN were used as statis- tical methods. For the first study (Luff et al. 1989), very heterogenous material from 248 sites in North-East England was used, primarily collected for use in the British mapping scheme. Ten habitat groups were recognized based on the distribution of the species presence over the sites. A method was developed to fit in new sites. A first limitation of this method is that new sites can only be fit in using species that already contributed to the orig- inal ordination. Secondly, that only pres- ence/absence data can be used. The original data set consists both of data from pitfalls, and sampling by hand. The possibility to include also samples from localities where pitfalls do not function, such as shore habitats, certainly is an advantage when using only the species presence. A main disadvan- tage is that the species’ relative occurrences cannot be taken into account. The study of Luff et al. is, as they also notice in their discussion, in the first place a methodological one and the results do not give a definitive and complete picture of carabid habitats of the British Isles. Many habitats, for instance woodland, will have to be investigated more inten- sively. Three of the ten habitats do not occur in The Netherlands (upland dry, upland wet and boulder). The remaining habitat groups are rather broadly defined: coastal, woodland, dry grassland, wet grassland, wet running, wet still - silt, wet still - sand. It is rather difficult to compare the species preferences for British habitats with the present results, although many species preferences seem to correlate well, e.g. those of Calathus erratus, Amara bifrons, Calathus rotundicollis and those of many eurytopic species such as Calathus melanocephalus, Loricera pilicornis, Pterostichus nigrita and Tre- chus obtusus. From the present classification it can be learned that the most important environmental factor influencing the ordination of carabid species is soil moisture (fig. 14). This is in support of the analyses by Luff et al. (1989). It is therefore very plausible that carabids are good indicators for moisture conditions in the field. In the study concerning the heathland and grass- land habitats in northern and central Europe, Eyre & Luff (1990a) used material from 638 heath and grassland pitfall sites. A number of these sites have also been used in the present study. Classification with TWINSPAN resulted in the identification of 17 habitat groups. In this study, woodland sites as well as shore habitats were not included. Although it is not easy to compare this classification with the present results, it is clear that some primary div- isions are similar, e.g. heath, cultivated land and polders (pioneer habitat sites). The classification of Eyre and Luff (1990b) con- cerns the classification of British grasslands. The data consist of material from all over England, Wales and Scotland. The results are to some extent supporting the present analysis. ‘Dry’ species, such as Broscus cephalotes, Calathus mollis and C. fus- cipes are found in the same habitats in Britain. Also heath and peat-moor species (group A, table 2) have been placed in similar habitat groups in the analysis of Eyre & Luff. Some species that are recognized as woodland species in the present anal- ysis, are scattered over many habitat groups in the British study. This is probably because mature woodlands have not been taken into account. In the results of both British studies (Eyre & Luff 1990b, Luff et al. 1989) some species preferences differ obviously from the situation in The Netherlands, e.g. the above mentioned species Carabus violaceus, Pterostichus cristatus and P. madidus seem to be much more eurytopic in England. In their discussion Eyre and Luff (1990a) men- tion the scale problem, stating that the greater the area covered by a study, the less precise the results will be. It is obvious that in such a case the classi- fication will be influenced by the geographical dis- tribution of the species. A geographical component is even present within a relatively small study area as The Netherlands, where, for instance, Pterosti- chus madidus occurs only in the southern and cen- tral part of The Netherlands. The mean In(SDY+1) in our method, however, is calculated over all samples of a certain habitat, also those situated outside the range of P. madidus. This prob- lem applies to all species with a restricted distribu- tion within a certain study area. The problem be- comes more pressing, however, when the study area is still larger, because the relative number of species with restricted distributions will increase. On the other hand too small areas such as the province of Drenthe in The Netherlands (Den Boer 1977) contain not enough habitats to make a classification useful. TURIN ET AL.: Carabid beetles The present results will probably surprise expe- rienced carabidologists to some extent. Several spe- cies show rather unexpected occurrences. Abax pa- rallelepipedus, for instance, is a species characte- rized by many authors as restricted to forests, forest edges and hedges (Koch 1989, Lindroth 1986, Thiele 1977, Turin & Heijerman 1988), but ap- pears to occur in a wide range of habitats abund- antly, and in an even wider range in low numbers. It is a well known fact that species preferences for certain types of vegetation (habitats) can shift over geographical distance or at different altitudes. Pterostichus cristatus, for instance, is an inhabitant of cool mountain Fagetalia forest (Thiele 1977), but it also occurs in open, moist sites in Cumber- land and Northumberland in the British Isles (Lin- droth 1974). Pterostichus madidus, which inhabits the Fagetalia and Querco-Carpinetum (oak-horn- beam forest) (Koch 1989, Thiele 1977) in Central Europe, prefers open country and cultivated soil in the British Isles (Lindroth 1974, Eyre & Luff 1990a, Luff et al. 1989). In The Netherlands it takes an intermediate position (table 5). In the southern part of the province of Limburg P. madidus is the most abundant species in the forest edges of the oak-hornbeam forest, but it also occurs quite fre- quently and abundantly in shaded meadows and limestone grasslands without an extremely south- ern exposition. In the central part of The Nether- lands it can only be found in light forest and in low numbers. The above mentioned differences be- tween the present classification and that of British Isles will partly be due to such geographical differ- ences in species habitat preferences. We support the conclusion of Eyre & Luff (1990a) that classifications based on large datasets, together with the data from pitfall sampling from all over Europe, make carabid beetles a reliable group for environmental monitoring. ACKNOWLEDGEMENTS We are grateful to the ‘Prins Bernhard Fonds’ for a grant enabling H. Turin to carry out this carabid project in the period 1982-1985. We also thank all institutions supporting the project: the Staatsbosbeheer, especially for providing the opportunity to collect the data and describe the sampling localities; the '"Biogeografisch In- formatie Centrum’ for a grant for computer processing of the data; The Dutch Central Office of European Inverte- brate Survey, the Netherlands Entomological Society, the Biological Station, Wijster, and especially the Research Institute for Natural Management, Arnhem, where we had our home for three years. The authors thank L. Brussaard, J. van Tol and an unknown referee for their valuable suggestions. We regret that our late colleague and co-author Eric Penterman is not able to see the final results of his work of building up a part of the database and especially uniformly describing and photographing all sampling localities. 289 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 REFERENCES Andersen, J. 1970. Habitat choice and life history of Bembidiini (Col. Carabidae) on river banks in central and northern Norway. - Norsk Entomologisk Tidssk- rift 17: 17-65. Andersen, J. 1982. Contribution to the knowledge of the distribution, habitat selection and life history of the riparian beetles in Norway. - Fauna norvegica (B) 29: 62-68. Andersen, J. 1983. The habitat distribution of species of the tribe Bembidiini (Coleoptera, Carabidae) on banks and shores in northern Norway. - Notulae Entomo- logicae. Helsingfors 63: 131-142. Aukema, B., 1990. Taxonomy, life-history and distribu- tion of three closely related species of the genus Ca- lathus Bonelli (Coleoptera: Carabidae). - Tijdschrift voor Entomologie 133: 121-141. Baars, M. A., 1979. Catches in pitfall traps in relation to mean densities of carabid beetles. - Oecologia (Berlin) 41: 25-46. Basedow, T., A. Borg, R. de Clercq., W. Nijveldt & E. Scherney, 1976. Untersuchungen tiber das Vorkom- men der Laufkäfer (Col.: Carabidae) auf europäischen Getreidefeldern. - Entomophaga 21 (1): 59-72. Becker, J., 1975. Art und Ursachen der Habitatbindung von Bodenarthropoden (Carabidae, Coleoptera, Di- plopoda, Isopoda) xerothermer Standorte in der Eifel. - Beiträge zur Landespflege, Rheinland-Pfalz, Beiheft 4: 89-140. Boer, P. J. den, 1965. Verbreitung von Carabiden und ihr Zusammenhang mit Vegetation und Boden. - In: R. Tüxen, Biosoziologie. Den Haag: 172-183. Boer, P. J. den, 1977. Dispersal power and survival. Ca- rabids in a cultivated countryside (with a mathemat- ical appendix by J. Reddingius) - Miscellaneous Pap- ers, Wageningen 14: 1-190. Boer, P. J. den, 1979. The individual behaviour and pop- ulation dynamics of some carabid beetles of forests. - In: P.J. den Boer et al., On the evolution of behaviour in carabid beetles. - Miscellaneous Papers, Wagenin- gen 18: 151-166. Boer, P. J. den, 1986. Concluding remarks. - In: P J. den Boer et al., Carabid beetles, their adaptations and dy- namics. — Fischer, Stuttgart, New York: 534-557. Boer, P. J. den, 1990. Density limits and survival of local populations in 64 carabid species with different pow- ers of dispersal. - Journal of evolutionary Biology 3: 19-48. David, J. & P. Marchal, 1963. Les coléopteres Carabiques du marais et Echets (Département de l’Ain): Contri- bution a l'étude d'un peuplement paludicole. - Bulletin mensuel de la Société Linnéenne de Lyon 32: 109-125. Dawson, N., 1965. A comparative study of the ecology of eight species of fenland Carabidae (Coleoptera). - Journal of Animal Ecology 34: 299-314. Desender K. & H. Turin, 1989. Loss of habitats and changes in the composition of the ground and tiger beetle fauna in four West European countries since 1950 (Coleoptera: Carabidae, Cicindelidae). - Biolog- ical Conservation 48: 277-294 Drift, J. van der, 1959. Field studies on the surface fauna of forests. - Bijdragen tot de dierkunde 29: 79-103. Eyre, M. D. & M. L. Luff, 1990a. A preliminary classifi- cation of european grassland habitats using carabid beetles. - In: N. E. Stork, The role of ground beetles 290 in ecological and environmental studies. - Intercept, Andover, Hampshire: 227-236. Eyre, M. D. & M. L. Luff, 1990b. The ground beetle (Coleoptera: Carabidae) assemblages of British grass- lands. - Entomologist’s Gazette 41: 197-208. Grossecappenberg, W., D. Mossakowski & F. Weber, 1978. Beitrage zur Kenntnis der terrestrischen Fauna des Gildenhauser Venns bei Bentheim. I. Die Carabi- denfauna der Heiden, Ufer und Moore. - Abhandlun- gen aus dem Landesmuseum für Naturkunde zu Münster in Westfalen 40: 12-34. Haeck, J., 1971. The immigration and settlement of Ca- rabidae in the new IJsselmeerpolders. - In: P. J. den Boer, Dispersal and dispersal power of carabid beetles. — Miscellaneous Papers, Wageningen 8: 33-52. Haeck, J., R. Hengeveld & H. Turin, 1980. Colonization of road verges in three Dutch polders by plants and ground beetles (Coleoptera: Carabidae). - Entomolo- gia Generalis 6: 201-215. Heckendorf, Chr., A. Ruprecht, K. Schneider & F. Tietze, 1986. Zur Faunenstruktur (Coleoptera-Carabidae) in Wald-Brachland-Habitaten des NSG 'Lintbusch’ (158). - Hercynia (Leipzig) N.E 23: 72-82. Heijerman, Th. & H. Turin. (1989). The carabid fauna of some types of forests in The Netherlands; a numerical comparative analysis. - Tijdschrift voor Entomologie 132: 241-250. Heydemann, B., 1962. Die biozönotische Entwicklung vom Vorland zum Koog, II Teil: Käfer (Coleoptera).- Abhandlungen der mathematisch-naturwissenschaf- tlichen Klasse der Akademie der Wissenschaften und Literatur, Mainz 11: 765-964. Heydemann, B., 1967. Die biologische Grenze Land-Meer im Bereich der Salzwiesen. - Steiner Verlag, Wies- baden: 200 pp. Hill, M. O., 1979a. TwINSPAN: A Fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. — Ithaca, New York. Hill, M. O., 1979b. DECORANA: A Fortran program for detrended correspondence analysis and reciprocal averaging. — Ithaca, New York. Horion, A. & K. Hoch, 1954. Beitrag zur Kenntnis der Koleopteren-Fauna der Rheinischen Moorgebiete. - Decheniana 102B: 9-39. Jarmer, G., 1971. Vergleich der Carabidenfauna des ‘Griether-buschs’ und des ‘Schwarzen Wassers’ am Niederrhein (eutrophe und dystrophe Standorte). - Jahresbericht der Universität Köln 1971: 290. Koch, D., 1984. Pterostichus nigrita, ein Komplex von Zwillingsarten. - Entomologische Blätter für Biologie und Systematik der Käfer 79: 141-152. Koch, K., 1977. Zur unterschiedlichen Besiedlung von Kiesgruben am Niederrhein duch ripicole käferarten. - Decheniana, Beihefte 20: 29-35. “à Koch, K., 1989. Die Käfer Mitteleuropas, Okologie Band 1.- In: H. Freude et al., Die Käfer Mitteleuropas, Band El. - Goecke & Evers, Krefeld: 1-107. Kolbe, W., 1968. Vergleich der bodenbewohnenden Co- leopteren aus zwei Eichen-Birken-Waldern. - Ento- mologische Zeitschrift 78: 140-144. Kolbe, W., 1970. Vergleichende Coleopterenfänge in zwei Siegerländer Laubwäldern. - Natur und Heimat 30: 22-256. Krogerus, H., 1948. Okologische Untersuchungen über Uferinsekten. - Acta Zoolologica Fennica 53: 1-157. Lauterbach, A. W., 1964. Verbreitungs- und aktivitàtsbes- timmente Faktoren bei Carabiden in sauerlandischen Waldern. - Abhandlungen aus dem Landesmuseum fiir Naturkunde zu Miinster in Westfalen 26: 1-103. Lehmann, H., 1965. Okologische Untersuchungen über die Carabidenfauna des Rheinufers in der Umgebung von Köln. - Zeitschrift für Morphologie und Ökologie der Tiere 55: 597-630. Lindroth, C. H., 1945. Die Fennoskandischen Carabiden I, Spezieller Teil. - Göteborg, 709 pp. Lindroth, C. H., 1949. Die Fennoskandischen Carabiden III, Algemeiner Teil. - Göteborg, 911 pp. Lindroth, C. H., 1974. Coleoptera, Carabidae. - Handbooks for the identification of British insects 4 (2): 1-148. Lindroth, C. H., 1985. The Carabidae (Coleoptera) of Fennoscandia and Denmark I. - Fauna Entomologica Scandinavica 15 (1): 1-226. Lindroth, C. H., 1986. The Carabidae (Coleoptera) of Fennoscandia and Denmark II. - Fauna Entomologica Scandinavica 15 (2): 227-497. Lohse, G. A., 1983. Die Asaphidion-Arten aus der Ver- wandtschaft des A. flavipes L. - Entomologische Blat- ter fiir Biologie und Systematik der Kafer 79: 33-36. Lompe, A., 1989. Erginzungen und Berichtigungen zu Freude-Harde-Lohse ‘Die Kafer Mitteleuropas, Band 2 Unterordnung Adephaga: 1, Familie Carabidae (1976)’ - Goecke & Evers, Krefeld: 23-59. Luff, M. L., 1975. Some features influencing the efficiency of pitfall traps. - Oecologia (Berlin) 19: 345-357. Luff, M. L., M. D. Eyre & S. P. Rushton, 1989. Classifica- tion and ordination of habitats of ground beetles (Co- leoptera, Carabidae) in north-east England. - Journal of Biogeography 16: 121-130. Meijer, J., 1973. Die Besiedlung des neuen Lauwerszee- polders durch Laufkäfer (Carabidae) und Spinnen (Aranea). — Faunistisch-ökologische Mitteilungen, Kiel 4: 169-184. Meijer, J., 1974. A comparative study of the immigration of carabids (Coleoptera, Carabidae) into a new polder. - Oecologia (Berlin) 16: 185-208. Meier, J., 1980. Colonization of the Lauwerszeepolder by some elements of the arthropod Fauna. — Disserta- tion, Vrije Universiteit, Amsterdam, 98 pp. Mossakowski, D. 1964. Uber Verbreitung und Okologie einiger Kifer in Mooren und Heiden Schleswig-Hol- steins (Coleoptera: Carabidae et Staphylinidae). - Faunistische Mitteilungen aus Norddeutschland 2: 106-111. Mossakowski, D., 1970a. Okologische Untersuchungen an epigäischen Coleopteren atlantischer Moor- und Heidestandorte. - Zeitschrift für Wissenschaftliche Zoologie, Leipzig 181: 233-316. Mossakowski, D., 1970b. Zur Besiedlung salzbeein- flussten Torf-Standorte durch Coleopteren. - Mittei- lungen der Deutschen bodenkundlichen Gesellschaft 10: 217-219. rs Mossakowski, D., 1970c. Das Hochmoor-Okareal von Agonum ericeti Panz. (Coleoptera, Carabidae) und die Frage der Hochmoorbindung. - Faunistisch-ökolo- gische Mitteilungen, Kiel 3: 378-392. Mossakowski, D. 1971. Okologische Untersuchungen der Coleopterenfauna salzbeeinflussten Torfe. - Verhand- lungen der naturwissenschaftlichen Heimatforschung 38: 117-131. Niemela, J., 1988. Carabid beetles in shore habitats on the Aland Islands, SW Finland: the effect of habitat avail- TURIN ET AL.: Carabid beetles ability and species characteristics. - Acta Oecologica, Oecologica Generalis 9: 379-395. Obrtel, R., 1972. Soil surface Coleoptera in a reed swamp. - Acta Scientiarum Naturalium Academia Scientia- rum Bohemoslovacae, Brno 6: 1-35. Penterman, E. & H. Turin, 1985. Handleiding en index bij de databank van de Nederlandse Loopkevergegev- ens. - European Invertebrate Survey-Nederland, Leiden, 94 pp. Plachter, H., 1986. Composition of the carabid fauna of natural riverbanks and of manmade secondary habi- tats. - In: P. J. den Boer et al., Carabid beetles, their adaptations and dynamics. — Fischer, Stuttgart, New York: 509-535. Rabeler, W., 1947. Die Tiergesellschaft der trockenen Calluna-heiden in Nordwestdeutschland. — Jahresbe- richt der Naturhistorischen Gesellschaft zu Hannover 94/98: 357-375. Rabeler, W., 1957. Die Tiergesellschaft eines Eichen- Birkenwaldes im nordwestdeutschen Altmoränenge- biet. - Mitteilungen der Flor. Soz. Arbeits Gemein- schaft 6/7: 297-319. Rabeler, W., 1962. Die Tiergesellschaft von Laubwäldern (Querco-Fageta) im oberen und mittleren Weserge- biet. - Mitteilungen der Flor. Soz. Arbeits Gemein- schaft, N. F. 9: 200-229. Rabeler, W., 1963. Charakterisierung der Streufauna ei- niger nordwestdeutscher Waldgesellschaften. - In: M. J. Doeksen & J. van der Drift. Soil organisms. - Ams- terdam. Rabeler, W., 1967. Zur Charakterisierung der Fi wald-Biozönose im Harz auf Grund der Spinnen- und Käferfauna. - Schriftenreihe für Vegetationskunde 2: 205-236. Rabeler, W., 1969. Zur Kenntnis des nordwestdeutschen Eichen-Birkenwaldfauna. - Schriftenreihe für Vegeta- tionskunde 4: 131-154. Renkonen, O., 1944. Die Carabiden und Staphyliniden- bestände eines Seeufers im SW-Finnland. — Annales Entomologici Fennici 10:. Rueda, F. & C. Montes, 1987. Riparian carabids of saline aquatic ecosystems. — Acta Phytopathologica et Ento- mologica Hungarica 22: 247-263. Schjotz-Christensen, B., 1957. The beetle fauna of the Corynephoretum in the ground of the Mols Labora- tory. - Natura Jutlandica 6/7: 1-20. Schjetz-Christensen, B., 1966a. Biology of some ground beetles (Harpalus Latr.) of the Corynephoretum. - Natura Jutlandica 12: 225-229. Schjetz-Christensen, B., 1966b. Some notes on the biol- ogy of Bradycellus collaris Payk. and B. similis Dej. (Col., Carabidae). - Natura Jutlandica 12: 230-234. Schweiger, H., 1975. Neue Asaphidion-Arten aus der Verwandtschaft des A.flavipes L. - Koleopterolo- gische Rundschau 52: 105-111. Simpson, E. H., 1949. Measurement of diversity. - Nature 163: 688. Szyszko, J., 1986. Dynamics of population size and devel- opment of the carabid fauna in pine stands of poor sandy soils (facts and suppositions). In: P. J. den Boer et al. Carabid beetles, their adaptations and dynamics. - Fischer, Stuttgart, New York: 331-341. Thiele, H. U., 1956. Die Tiergesellschaften der Boden- streu in den verschiedenen Waldtypen des Niederber- gischen Landes. - Zeitschrift für angewandte Ento- mologie 39: 316-357. 291 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Thiele, H. U., 1977. Carabid beetles in their environ- ments. A study on habitat selection by adaptions in physiology and behaviour. - Zoophysiology an Ecol- ogy 10. Springer, Berlin, Heidelberg, New York: I- XVII, 1-369. Tol, J. van, 1981. Handieiding en atlas voor het mede- werken aan entomologische projekten in het kader van de European Invertebrate Survey. - Leiden: 1-60, maps. Turin, H., 1983. De invertebratenfauna van de Zuidlim- burgse kalkgraslanden. Loopkevers van de kalkgras- landen en hellingbossen. — Natuurhistorisch Maand- blad 72: 72-83. Turin, H., 1990. Naamlijst voor de Nederlandse loopkev- ers (Coleoptera: Carabidae). - Entomologische Be- richten, Amsterdam 50: 61-72. Turin, H., 1990. De loopkevers van het Waddengebied. - Entomologische Berichten, Amsterdam 51: 69-78. Turin, H., J. Haeck & R. Hengeveld, 1977. Atlas of the carabid beetles of The Netherlands. — Koninklijke TABLES Nederlandse Akademie van Wetenschappen, Verhan- delingen Afdeling Natuurkunde, Tweede reeks 68, Amsterdam, 228 pp. Turin, H. & Th. Heijerman, 1988. Ecological classifica- tion of forest-dwelling Carabidae (Coleoptera) in The Netherlands. - Tijdschrift voor Entomologie 131: 65- zi Turin, H. & E. Penterman, 1985. Dertig jaar loopkever- onderzoek met vangpotten. - Nieuwsbrief European Invertebrate Survey-Nederland 16: 35-46. Wasner, U., 1977. Die Exrophilus-Arten (Agonum, Cara- bidae, Coleoptera) des Federseerieds. Vergleichende Studien zur Okologie sympatrischer Arten engster Verwandtschaft. - Thesis, Tübingen: 1-316. Received: 8 July 1991 Revised version accepted: 1 November 1991 Table 1. The 33 habitats recognized by the ‘ecocode’, and per habitat, the numbers of year-samples and the number of sites where pitfall series were situated. NR HABITAT 1 Peat moor (wet, Sphagnum) 2 Heath with Molinia (moist) 3 Erica heath (moist) 4 Calluna heath (dry) 5 Heath with grasses, Deschampsia (dry) 6 Corynethoretum (dry, open) 7 Coastal dunes (dry, open) 8 Dune grassland (coastal) 9 Dune forest (coastal) 10 Dune scrub (coastal) 11 Poor grassland on sandy soil (inland) 12 Cultivated, pasture (inland, sand, fertilized) 13 Cultivated, arable (inland, sand, fertilized) 14 Cultivated, waste land (inland, sand) 15 Coniferous forest, plantation (open) 16 Coniferous forest, mature 17 Coniferous forest, old (moist) 18 Deciduous forest, oak-birch 19 Deciduous forest, oak-beech 20 Deciduous forest, oak-hornbeam 21 Deciduous forest, poplar (moist, polders) 22 Deciduous forest, alder-willow (moist-wet) 23 Inland scrub (moist) 24 Ruderal, park, orchard (dynamic) 25 Limestone grassland, dike slopes (xerotherm) 26 Grassland with herbs (inland, unfertilized) 27 Reedland, Lauwersmeer polder (sand-silt) 28 Reedland, IJsselmeer polder (sea clay) 29 Cultivated, IJsselmeer polder (sea clay) 30 Colonization sites, building lands (recent) 31 Sand banks near salt water, seashore (open) 32 Inland shores, river banks (open) 33 Salt marshes (sea clay) SAMPLES SITES n n 16 9 28 9 14 7 101 27 119 26 65 24 53 20 69 29 68 13 100 >Il 23 14 33 22 17 17 26 11 15 12 65 64 11 11 126 73 41 34 17 17 19 13 42 40 44 22 40 40 54 54 98 70 18 4 D? 22 52 26 35 15 93 40 22 6 40 20 Tables 2-9. The tables show the distribution of species relative occurrences over the 33 habitats recognized. Explanation A-H. - Tables 2-7, show the main species groups A-H, recognized by TWINSPAN classification. The sub-division of the main species-groups B and D is based on a sixth level, and those of groups E, F, G and H is based on a fourth level TWINSPAN division. EU, RA. - The tables 8 and 9 show respectively the eurotopic (EU) and rare (RA) species, which have been separated from the main groups A-H. Eurytopic species (Pres > 0.75 or Sim > 0.85), have been placed into one group (table 8) to get a more clear picture of the most typical species of the principal groups A-H. The rare species have been separated from these groups for statis- tical reasons. They occur in too low numbers in The Netherlands (Sa < 6 and N < 50, except when all were found in the same habitat), to get a reliable picture of the species habitat preferences NUM. - Species numbers. The full names with refer- ences to this number, can be found in the index GR. - In tables 2-7 in this column the species group subdivision is indicated. In tables 8 and 9 the letters A- H refer to the original primary group where the eurytopic and rare species were placed by TWINSPAN classification I-VII. - In the heading of the tables: main habitat groups that have been recognized by TWINSPAN classifi- cation (see fig. 4). 1-33. - In the heading of the tables: habitats, explana- tion see table 1. TURIN ET AL.: Carabid beetles 1-9 and *. - In the body of the table, give relative occurrences according to the percentual rescaling per species (see text): ‘ = In(SDY+1) <5%, ‘1 = 5% < In(SDY+1) < 15%, 2’ = 15% 2, otherwise no indication of soil preference is given: c = riverclay + seaclay, li = limestone, Il = limestone + loam, lo = loam, ls = loamy sand / sandy clay, pm = peatmoor, rc = river clay, sa = sand, sc = sea clay, sm = sand + peatmoor. Hu. - Humidity-preference measure (dry 456 samples, moist 690 samples, wet 461 samples), based on mean In(SDY+1). 1 = mainly in dry samples [dry > 10 X (moist+wet)], 2 = not in wet samples [dry+moist > 20 X (wet)], 3 = not in dry and not in wet samples [moist > 10 X (dry+ wet) ], 4 = not in dry samples [wet+moist > 20 X (dry)], 5 = mainly in wet samples [wet > 10 X (dry+moist) | Sa. - Number of year-samples N. - Number of specimens. 293 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 OCT LS ST OT % VI £06 801 OFT 08 pOTOT 087 MINA es es es (74 (19 SS 89° UL v8 OS WIS Saig | CE ZE TE|OE 67 87 LL 9T|ST HC EL U IC IA A Sv Id cl 1d sr 1d 0¢ Id vg * D) wl Id Ly 9 Ce SIS i Se 8 1d waaasnfanaafno 6s ONE MON MIRO) SEE Pe. bate Bie. EEE PIIMI eJewy smojids snrwoig BUISTISOUOU SEIMILA] eplaqdy epapunr) smernsqns snjrydonoN sandıqwe snyieye) sanadswnN T4 ‘14 dno13 sanads 0% 67 87 LT 9% ST 'suoneJuejd snozajruoo pue puer spgese Ápues Jo sarads :7g ‘pue[ssesd 100d pur ‘syexqeu aunp ‘4749409 daul409 30 sanads :[ gq ‘uont1adaa asıeds qu spros Ápues srydono#ro-uou zo samadg ‘€ aqeL LG LI 6 OU + die ot es SI EL wd EEC ZL ws LI CIC zese O8T wd ZI L wd 00p Em wd Phys SLT wd PRZEI LOE wd LC “GUI wd 9 € ¢ wd 6 9 y wd 61 9 SI TBL 66 wd pr val BT SL wd 6681 ZLI ws GGIT 16 wd 18 y ¢ ud OSLI HOT wd CON S8Ir ch L ¢ wd 08° OL 09° cL SL v8 95 68 £8 8L 08 09" 00° 0° 19° 85° So 6L SS OL 0° SL C8’ 87° kl NRE SE OS UI SHS ARC CNE NO CSC SC; IL GO TACO EZ TT AT NORG TER TAI OTC UI CISTI [072 6 Bh UR = en TND Br vg TRI i i II I . el vl È à IRE Ege Ae EV; po | z I i ee Aat eN IEP 3 ; B Aen ST G x| 6 à IV 0€ * I I Gl) Ge RY 6€ I * Cla 6 ZEV: OL Lel (6 I ÿ 4 + STARE CAV SLY 0¢ ; ; e e SW SA 60° 4 G * IV 9 j I ae ARE y Pe Or Ge Oy cS ae i ali I ‘ ue ¢ re Si BN er Ge AA 6€ ; : ; x SERE VARA as ATEN IV 9 È i Gi ‘ È VAI SER IV Co * IV 90° * Ok IV ST G 14 (SIN IC > i iz En Gol IV (GI C {8% IV FT 0 I i (AEREE E ee NY 0€ al j ; | [x x IV 174 e 60° «| IV SA I 3 I RSI: : : I i A ERI Cis “AI ay gl “I ; sam 1 CNE > : : i o È a i SA Aje ABe al ANY cls y IV winjequndxas wnuody JOUTW SNYMSOIIIY eWIJUI PIELV sınsadwe) eJapunT) s11exjos snjedieH smej snjedseH smeudo9 snjjaooy sy, suenyisae snpiydonon SMIE][IIULI SOGLIE) sıjos1yns snqjaodperg snpidaj sny2ns01314 Smepumo snydo1si]o sndeariowiau SN|AIEPOSIUY myprudıy unuody wninsiaA wnuody wnsoresodeA sipurwf9) PIMDIE LIOPOISTI suaiu SOLI) sinsanba eıewy nadia wnuody SNUIII9IE SNYNISOIIIT SISUDAIE SngeIe) suadipp sSNUINSOISIA snignp sndjednoy mm ON anna oO N AN OT U O0 M™ ODO WA A HNO N ODN n Hi Hun sanadswnN LV dnosd sarads ‘sos Apues arydonoërjo 10/pue sqros Ayead Jo sarads ‘7 o[geL 1661 ‘PET INNTOA ‘AIDOIOWOLNY YOOA LaraHosafiy, TURIN ET AL.: Carabid beetles 1644074 6LL TOI EVI SSI ESSI 668 6l 9811 107 PRI 67 107 CSCI 9L9 18€ LE CIE 957 174 067 6b cL 9% 656 901 LOST TIL TI €59 188 Ops 198} Lp NN DAMN N OD anna Nm ON NO MN NAN et et ed 4 es WS ES ol es ws es BS es ws es US es ES es es es ws es wd es es ES ES es I N ES NH OS WIS Said ice ze IE (4S 19° ST cy 66 Colin en ; * Cam yl CIC : z Ge A HE (RE HANSE yo AG WANG OL 281 VAL NCIC AD AGC PALIT WC (eh ee OI Ges 1G SI][ow snyseje> PANY eleWy I]asuanb eıewy SIJE][09 PUTATTD sijeiqn elewy 19 dn018 sanadg sapadswnN 'samads-g aya JO asoYI WO 1ajzrp ivy) ‘says uado wo samadg ‘7 Arge] N — Gal en * . - CNNOL RE on = man — — N N EGNA N VO al n n n IE GN eta — neo * * a 1d smejoaaozrIpenb ‘1501914 snı3Jdonsıw snyzeje sidjedijns snjedieH 14114201} snjedieH SMIIUII SOYIE]EI) snuipdezews snjedseH snixue snjedieH eosny view y STJE|NSUOI eIeWy eUIJeS eIIgaN siqseqruids sn3sta] e[091119) SNUIISOWIIL] snpuandunsıp snjedseH sajojeydad snosorg eigoysue eiewy EIIEAJÁS EJ9PUNT) sn199[Fau snjedseH Smep102 snuoydo e30UIJÁ9 vIeWY sıuuadıaıd snjedieH epiony LILLY sijezawny SIPUIUAT) TÁurwa8 snyrydonoN s1ejnsew SIpuru AT) AUIODHISIU UOIPIQUEY I[EYINIA SNIIOSE sndoyuex snjedseH SIjeuJaA snjedseH snAJ9s snjedseH smepmsndiq snaedeueg Gy 42 Cp Ty Ih Ov 6€ 8 Le 9€ Se pe ce ce I¢ ‘(panunuos) ‘€ olde L 295 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 LE LI [4 98851 19€ 87 Ic [4 £9 9 € ISPE 907 I 67 y CO IE [4 GI 8 [4 ly 9 £ LI6T y € 070% PLE L607 BEI 2 OZLGT LT GLLI $8 y 61h 907 LEE 88 [4 IST SS [4 LIIp 08 [ 001 LL lè col SS G ey 9 y 61 9 y €01 PI £ Kl HB LELT 821 y 2 G 96 9 [4 9 y [ TESTS c DI y [4 69 8 (È SL £ I GEL CZ C 169 06 [4 Ia) 4 [4 87 € I £8 L 14 87 ST NC GW 14 v0 8L 69 24 00° Ly’ 19° uns Said ; 5 I EEE NEE SEELTEIU SEE VA EO A WANNA OE U Clee Ol IIA enni 8 Ee Ban, ST pl I I I el I S VEG 1) a9 a ae ‘(çQ) s1S910J 10 ‘(Z7q) [ros Ayyeyd UO saas papeys 3srow ‘(T(]) SPueIsse1d JuoIsaun] 10 [eIspnI ‘pareArajno zayare ‘sais papeys woa saroads ‘G 3]qeL Gil WU OL GERS TO) essiwajaeid erewy snoneuajgoud sngeıe) sı1de snrworq 101 1937 UOIPIQUISY 001 SNILUITILWOJNI SMISIOT 66 stunuwos0pnasd erewy 86 eauunig eıewy /6 SUINU SNUYPNOYNIT, 96 snaoid sdojow 66 sojajjesed xeqy 46 smenpundoduo]go 1501314 CG SNaDEJOIA SOGLIE) 76 snpadidajayjesed xeqy 16 smepundrupenb snjedseH 06 sadyni snrydonon 68 saploqered snIYDAY gg ol TOL snjeine snqeie) /8 snpıpew snyoI3S0191d 98 sijruow snqeie) Cg SMIILIIOI SOGLIE) $8 ıdıeys snjjaskpeig Cg suaal] wnuo8y 78 sijedas smydedg 18 SIAE GIA SBE 08 smesrwnd SOS GL e|eyda0010]Y9 eig] 8/ na]pw snuoydQ // smerprwip snjedieH 9/ Suey1d21 snuryseig G/ epnmu LILY #/ sadiapi3 vorprquag €/ sluzooijnsew snuoydoreg Z/ sIqregigni snuoydo IL sadisqni snpedieH 0/ sdanound snuoydg 69 smenuane snjedieH 89 ESEANUOWI eıewy /9 210818 umuody 99 sIPIASNAN ¢q-1q dno:8 sanads 296 TURIN ET AL.: Carabid beetles lai Ee LST T nH ST 68 DICI, LE DI gg 31 18; INC: ud 6/ I 199; 18° OSS el ce 19° 60° ST IT sr 0€ PT 9 sr ST cy SS 0¢ sr cl 42 IT SS CL 44 Cla IT 1G; Id LE 44 Sold |= al n GE Ae NG (e ©) teri SERA À toni * END SIE Ee CO ESE Ei EBEN | ae EAU) LILLY TC] ıolewxnı snaedeueg (I wnıssow wnuody (GI snsourdipn snayderg 8ZI sapropedsey vorpiquiag [ZI snumesysue SNYNSOIIIT OZ] wnyepundıwas vorpiqwag GZI LUNI] [SUS PUOI PI AUS TT CT wmenndiq vorpiqwag €71 PIPAO eIewy ZZ] umequi] uosydowg 171 sajpdesine sernawag OZI winsnjqo uoIpiquiag GII smemurid sngese) SI saprordopay sapoQ LIT sıppooraegj sndjedmy II ejooidna snuoydQ SII snynione snqoned II STÉPOS MONTE a] I9SEQIA snyjaokpesg ZII sapperd SOL III sadi][ed uorprydesy QT] suoijiq erewy 601 smejmewspenb sniwosiq 801 sıpeuorprzaus Jaisipeg LOT smejmsndrun ssısıpeg 901 snueipiiaw sndjedmy SOI wnnpra umuody HOI sanadswnN 49-15 ‘TA- LA ‘T4 dno1d sanadg (ED TDT) salads pare[aI pue (79) sMopeaw 1aM pue saysiewi (75) puejp931 ‘(ZA TA) sans papeys er jo Ajurew ‘sanads snoaue]|2SIN ‘9 2192 L 297 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 co n= quan (©) xt — DTD DD st \O nn + ST 2S SI 3s 2S 3s gs 00° <0 TS oy 00° LS 99° cy 09° IS ES Ly BL BL 69 18 cL’ 18° 08 vl 85 69 LIA 8L 65 NG 09; LT vl I SE po vl 0° SL cl IL or 8° 19 GL 18 WIS N — \O UNISER RICE, Usi N (ca FIA NOTATO TIENE ul N ila e OORUNI EEEN EON EE EEE A * * Sq cc ce IE OX BOTAN AON OD * GQ* NANNA N COENEN EK O Tei ATTIRARE UNE OMG GS e VONDA OU HE OESO FAL Gl ll WE OL G WO eG UD) aaa i i auvadipI]]ed vorpiqwag €41 snyyunsıp snyjaokpeig Z/I snjgjosqo snYINONIYIA {LT XO PA UO PI QUIS ZAT sruuadrpianj snuodog 691 WNUUEUIIOU UCIPIQUIg 89] snasjeyd snuodog [91 wnjoausdıe UOIpIquiag 99] STIDEIOY] snisiysskq SOT VAN LASHER FO sninosqo sniaruosÁ COT snuijes SNA 791 TARISNG SNYNINONIYN(K [9] COCO UOT pu ROD] wnause UOIPiquag 661 wumeudiew wnuody gC] winjeundigq vorpiquisg /ST snypod snruryoskqQ IST wngepnyewispenb uorpiqwag CCI P[NISNIXIAUOD PILUY HCI wnzezowaj voIpiqwag CGI snjnased sndjedmy 76] snredia sniydeyq TSI wmejnunj vorpıqwag OCT snaidno snzuderg GI Isapany snınydsÄqQ gy] LONGER 45E wnedrung UOIPIQU OFT [NESS WOT RIEL, Gil 1f910Y]) umuody Fh] CNT ENS ejnosnfew eıewy Zp] sayxıw snydojouaig TH] sadiqe wnuody Op] SSU PXI OPTA HEGEL SIUIONISIU SNIUAEJUT) ZEI SnosIp SNYPINOISET /¢] snngixa sndjednoy 96] SMEJOUIQ snjÁpepostuy GE] PINE eıewy HE] snaidnd sny9nsossıg CCI PIES eıewy ZC] saadsWAN PH-1H dno13 saisadg (PH) siearqey aroys Surpnjour ‘(CH-TH) Sergey 39M 03 IsTOW Jo sardads 1aau01q ‘/ [TEL 298 TURIN ET AL.: Carabid beetles ws ws ws uis ES wd IL 69 uns 001 Sad n en Ta * ÿ 9 4 ì I EMISE Bez re ù I CY. GG * i I EG Ay Gl I Is e ali TNT I 6. Go Oo 3G I 4 MENT EL ee CO ene SC EC El i PIV le: Be t eee © [4 I G I € x fe È ¢ y € O (Gee AT 14 È ¢ I I GG I à if m — n * O * 08 6G 822 [29T Ce CNT el I G. G I I If Ue Sl GC | _£ 6 PG Gs Ae eel Ug GC Ape I i Be ae dl es PA TT I mae ec È (Gi ea Ia Gli Os I IG [4 el GIG EZIO = bh AP 37 6 Ti Ale TEE ae WS * CC I oll DRE GS hE I [4 ¢ ¥ 8. 1 I ; € LG a pal I bp 9 « y i Tesi CE TG al T eT ere eG v8 el 2e de SG Il a | licence [4 eG rd [4 TE CE i I AGAINST I I oe I b y I i I ; We Ge TI Gee (Lion ees Cee Ory ; I (Comm loose TESTI I ne Cr a, I I I I i 61 x biG » Ç « Ih [ee ell I Cr ST] PS Le VARE Cor | nee Le € x 6 CT] Se I I I Be ce i CORTE ACen Os Carel ye Ge ie ¢ CE pb: DARAI [4 i I I I Le aS SZ LOT ayı U29MI9q UOSTIedUIOD Y ‘OT AEL I TER 00 £0" * H snaojeya snıny9siq 687 14 5 00 €0' * H 2[219)P] vorpiqwag 87 AE az OG LB AW ee: ; H snpriu sniryosÁq €87 [4 Cama GR SWE I H wanbijgo uorpiquiag 787 14 [4 90; 9 * H suedaja sndjednay 187 [4 7 Ob 90 VA REI H wmegnsndipenb uorpiqwag 087 ld 7 CE 290; * È H wmzeuny vorpiqwag 6/7 CI Ge 8 N * H snuesyswrsys snydojouaig 8/7 I I ¢ 00° €0 * H SISI) SNIUIEIUD L/T 8 Sr ee GEO); ¢ * Cl H PINUE]ILI EYUMEPO 9/7 L € ve 90 € * H sadiuuniq sndjednoy 6/7 T BEE 00 €0' * H snjewoue Jaisipeg H/Z ¢ € 00 €0 * H smandsuoo sndjednoy ¢/7 9 SIA SI L * H Snauae SNIHUISAG 7/7 L Ge II ay L * I H SNJIS2A smiuaeyyy 1/7 9 € IE 90) * 7 H suaqni snyaıL (027 € CAT, 65 60° T 9 xl 9 smei]ad zorsipeg 697 ÿ Cat (Se 90) * 6 5) snası13 snjedieH 897 T GE OSS) 8| x 5 snamnze snuoydQ /97 6 ¢ co ST *| x CA STAT 5 Hrs) snyfarkperg 997 € ee) € * 7 SMeIe Ip JOISIPEE COZ y PR Ce Slee’ Some 60 ¢ ARE I wnayıd wnuosy Hoz 7 T 05 90° 6 * q ewISIS sMIWOIG €97 (Ti e BR ZOO) 7 * q SUBI wnuody 797 y ¢ I 6y 90 * 8 a enuans eıewvy 197 Qi 77 € € 60 I * d SNINEUI SIISIOINY 09T [4 GI I ve 90 7 * d snisauoy snjedieH 657 I [ae xt 00 £0" * d snaeudisiipenb sniwoiq 867 I Te ee 00 £0 * a sije18 snyonsomrd LGL ‘(panunuo)) ‘6 olde L 301 TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Alphabetic list of species names with index to tables 2-9 Abbreviations: NUM = species number (table 2-9); So = soil preference; Hu = humidity preference; GR = species group; T = table number. For explanation see table 2-9, pp. 293-301. Species Abax carinatus (Duftschmid) Abax parallelepipedus (Piller & Mitterpacher)............. Abax parallelus (Duftschmid) Acupalpus brunnipes (Sturm) Acupalpus consputus (Duftschmid)................ Acupalpus dubius Schilsky..... Acupalpus elegans (Dejean) Acupalpus exiguus (Dejean.... Acupalpus flavicollis (Sturm) Acupalpus meridianus (L.)..... Acupalpus parvulus (Sturm) Agonum albipes (F.)............ Agonum assimile (Paykull).... Agonum dorsale (Pontoppi- dam) ee Se Agonum ericeti (Panzer)....... Agonum fuliginosum (PANZEL) en Agonum gracile (Gyllenhal) Agonum krynickii (Sperk)..... Agonum livens (Gyllenhal).... Agonum marginatum (L.)..... Agonum micans (Nicolai)..... Agonum moestum (Duftsch- MILE n Agonum muelleri (Herbst) … Agonum munsteri (Hellen) … Agonum obscurum (Herbst) Agonum piceum (L.)........... Agonum sexpunctatum (L.)... Agonum thoreyi Dejean....... Agonum versutum Sturm...... Agonum viduum (Panzer)..... Amara aenea (Degeer)......... Amara anthobia A. Villa & GBA ER te Amara apricaria (Paykull)..... Amara aulica (Panzer)......... Amara bifrons (Gyllenhal)..... Amara brunnea (Gyllenhal)... Amara communis (Panzer).... Amara consularis (Duftsch- DMI Amara convexior Stephens.... Amara convexiuscula (Mar- NAM mera ene MII ee Amara curta Dejean............ Amara equestris (Duftsch- MOLTA Amara eyrinota (Panzer)...... Amara famelica Zimmer- Amara familiaris (Duftsch- MIC A RI. Amara fulva (Mueller)......... Amara fusca Dejean............ 245 91 94 275 273 1 281 136 116 Il Il pm sc Is li Is Is pm pm pm rc Is rc Is Is Sa IC sa sa NUM So Hu 5 AN N rN N EN A N Nu GR RA D3 D3 RA RA Al RA H2 G2 El H2 H2 EU EU Al EU D1 Al D2 H4 RA G4 EU RA EU RA Al H2 Al El EU B2 EU H2 F1 D3 EU B2 EU H3 RA Bl Al Bl G4 EU Gil B2 COMNNNNO 00 \D COD VO NWN LA O0 N 00 nN DN WW \O N 00 WW CON OAN CW WW À 00 Species Amara infima (Duftschmid) Amara lucida (Duftschmid).... Amara lunicollis Schioedte..... Amara majuscula Chaudoir.... Amara montivaga Sturm....... Amara nitida Sturm............ Amara ovata (E)... Amara plebeja (Gyllenhal)..... Amara praetermissa (C.R. Salil berg) RN Amara pseudocommunis Bu- rakowski e EEA ee Amara quenseli (Schoenherr) Amara similata (Gyllenhal).... Amara spreta Dejean........... Amara strenua Zimmermann Amara tibialis (Paykull)........ Anisodactylus binotatus (F.) Anisodactylus nemorivagus @uftschmid) esse Asaphidion flavipes (L.)....... Asaphidion pallipes (Duftsch- MELA Badister anomalus (Perris).... Badister bullatus (Schrank).... Badister dilatatus Chaudoir.... Badister lacertosus Sturm...... Badister meridionalis Puel..... Badister peltatus (Panzer)..... Badister sodalis (Duftschmid) Badister unipustulatus Bonelli Bembidion aeneum Germar... Bembidion argenteolum ANAS eee Bembidion assimile Gyllenhal Bembidion biguttatum (F.)..... Bembidion bipunctatum (L.) Bembidion bruxellense Wes- Bembidion deletum Serville... Bembidion dentelleum (hun Dero) Mee Bembidion doris (Panzer)... Bembidion femoratum Sturm Bembidion fumigatum (Duftschmid) nnn Bembidion genei Kuester...... Bembidion gilvipes Sturm..... Bembidion guttula (F.)......... Bembidion harpaloides Ser- VIE RASE Bembidion humerale Sturm... Bembidion iricolor Bedel....... Bembidion lampros (Herbst) Bembidion laterale (Sa- mouelle) ees tere eee Bembidion lunatum (Duftsch- TA) E ER Bembidion lunulatum (Four- ELOY) in ne EE Tee: So Hu fa 2 ga 2 SC n 2 lo 2 Ii 2 pm 4 c 1 pm 5 Iki 2 3 pm 5 rc 4 pm 5 5 rc Ih dl me. di 5 SC Toms TGZ Cc pm 5 Sc 75) 5 4 GR Al Bl EU H2 DI DI G4 EU D3 D3 ei Hl EU RA Cl H2 Al EU F1 RA EU RA EU El RA F2 El H4 H4 H2 G4 H4 H2 RA G4 RA H3 H2 D3 D1 EU G4 RA H2 EU RA RA H2 Do} Do) oo oo oo LA LA —J NOD OS SANS NI WOO WOOD DN HA Ÿ OO SA LA Nn OO © LA LA N COW LN 4 N Species Bembidion mannerheimi C.R. Sahlberg RR e eer Bembidion minimum (F.)...... Bembidion nigricorne Gyllen- a end Bembidion obliquum Sturm... Bembidion obtusum Serville Bembidion pallidipenne (Il- Eee Bembidion properans (Ste- phens) en. worse seco. Bembidion quadrimaculatum (eee scan. Bembidion quadripustulatum SERVICE eenen Bembidion quinquestriatum Gyllenhal SMA een. Bembidion semipunctatum (Donovan) sheen. aes one Bembidion tetracolum Say..... Bembidion varium (Olivier) Bembidion velox (L.)........... Blethisa multipunctata (L.).... Brachinus crepitans (L.)........ Bradycellus caucasicus (Chau- HOUD) er er TDI Bradycellus csikii Laczo......... Bradycellus distinctus (De- jean) Bradycellus harpalinus (Ser- VIE) RR Bradycellus ruficollis (Ste- PRES) EAN ES ae Bradycellus sharpi Joy.......... Bradycellus verbasci (Duftsch- ee san: Broscus cephalotes (L.)......... Calathus ambiguus (Paykull) Calathus cinctus Motschulsky Calathus erratus (C.R. Sahl- Der Calathus fuscipes (Goeze)...... Calathus melanocephalus (L.) Calathus micropterus (Duftschmid)................ Calathus mollis (Marsham).... Calathus rotundicollis Dejean Calosoma inquisitor (L.)....... Carabus arvensis Herbst....... Carabus auratus L............... Carabus auronitens F............ Carabus cancellatus Illiger..... Carabus clathratus L............ Carabus convexus F............. Carabus coriaceus L............. Carabus granulatus L............ Carabus monilis F............... Carabus nemoralis Mueller.... Carabus nitens L................. Carabus problematicus Herbst Carabus violaceus L............. Chlaenius nigricornis (F.)...... Chlaenius tristis (Schaller)..... Chlaenius vestitus (Paykull) So Hu pm 5 Bd pe? 4 5 li SAMO me 5) rc Is sa 5 later? sm sa 4 sm pm fo’ 2 sa sm 2 sm sm Ge 102 sa 3 pm [10007 lowe pm 5 It 2 I 2 Mo 2 pm Il 2 5 me 9 GR RA H4 BI H4 RA G3 H4 EU H3 RA RA G4 EU H4 H4 RA DI EU RA H4 EU Al D2 F2 B2 Bl B2 EU EU EU B2 Cl EU RA Al D2 RA Al RA RA D2 G2 D2 EU Al D3 D3 H2 RA RA Nn \O 00 NI ANNA \O \O oo 00 OO WW DW DN ON A LA ND OO LA EN LA O ON NY \O LA ND KO 00 HW TURIN ET AL.: Carabid beetles Species Cicindela campestris L.......... Cicindela germanica L.......... Cicindela hybrida L.............. Cicindela maritima Latreille BANDE) CANTI E tt Cicindela sylvatica L............ Clivina collaris (Herbst)....... Clivina fossor (L.)............... Cychrus caraboides (L.)......... Cymindis humeralis (Four- CLONI TONI ed Cymindis macularis Fischer von Waldheim............... Cymindis vaporariorum (L.) Demetrias atricapillus (L.)..... Demetrias monostigma Sa- mouelle EE ME Le Erotche. Ates gn Dicheirotrichus obsoletus (Dejean) AMER Ae es Dromius agilis (F.).............- Dromius angustus Brulle....... Dromius linearis (Olivier)..... Dromius melanocephalus De- JARRE Dromius quadrimaculatus (L.) Dromius quadrisignatus (De- JR o Dromius sigma (Rossi)........ Dromius spilotus (Illiger)...... Dyschirius aeneus (Dejean).... Dyschirius angustatus (Ahrens) nn Nr Dyschirius chalceus Erichson Dyschirius globosus (Herbst) Dyschirius luedersi Wagner.... Dyschirius nitidus (Dejean)... Dyschirius obscurus (Gyllen- Hal ee Dyschirius politus (Dejean)... Dyschirius salinus Schaum..... Dyschirius thoracicus (Rossi) Elaphrus cupreus Duftschmid Elaphrus riparius (L.).......... Epaphius secalis (Paykull)..... Harpalus affinis (Schrank)..... Harpalus anxius (Duftsch- IMI) LT Harpalus attenuatus Stephens Harpalus calceatus (Duftsch- TTI) ERO RENEE NN tee Harpalus dimidiatus ROSSE Harpalus distinguendus (Buftschmid) ARE n Harpalus flavescens (Piller & Mitterpachen) Karren. Harpalus froehlichi Sturm..... Harpalus griseus (Panzer)..... Harpalus honestus (Duftsch- MIETTA Harpalus latus (L.).............. Harpalus luteicornis (Duftschmid)................ NUM SoHu 21 pm 229 27 1 239 45 62 205 88 lo 2 AO sa 2 Sa san] 9 pm 120 li 2. Casa: OL ISS 17,100 RZ 1018107 22 232 1 196 2 217 108 rc 258 3 263 29 2 272 ige 105 DAD hl 285 5 206 148 Is 283. ASCA 163 5 156 162s Isa 165 149 Is 5 151 128 Is el de 5 213 DAS GE nl 250 76 IN ASNSA Me? 237 1 U BA 3 268 3 259 1 19 234 2 NN vw Ww WN 00 „a Us \D TOO. Woe ONAN N NN I \O MI 00 \O \O \O WOO G 00 oo \O LA No DI \O TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991 Species Harpalus melancholicus De- EA ce Re Harpalus neglectus Serville.... Harpalus picipennis (Buftschmid) an... Harpalus quadripunctatus De- Ja dea Harpalus rubripes (Duftsch- ITV) AR EG MP RUE Un Harpalus rufipalpis Sturm..... Harpalus rufipes (Degeer)..... Harpalus serripes (Quensel) Harpalus servus (Duftsch- mid) tse ERAAN SERIE ah, Harpalus smaragdinus (Duftschmid) … … ae Harpalus solitaris Dejean...... Harpalus tardus (Panzer) … … Harpalus vernalis (Duftsch- TIA) EE Harpalus xanthopus Gem- miger & Harold............. Laemostenus terricola (FICIDSO mn Lasiotrechus discus (F.)......... Lebia chlorocephala (Hof- fr AT) E e ER Lebia cruxminor (L.)........... Leistus ferrugineus (L.)........ Leistus fulvibarbis Dejean..... Leistus rufomarginatus (Duftschmid)................ Leistus spinibarbis (F.)......... Leistus terminatus (Hellwig) Loricera pilicornis (F.).......... Masoreus wetterhali (Gyllen- ELISA u ns Microlestes maurus (Sturm) Microlestes minutulus (Goeze) an n ne Miscodera arctica (Paykull).... Molops piceus (Panzer)........ Nebria brevicollis (F.).......... Nebria livida (L.)............... Nebria salina Fairmaire & La- boulbene AREA te Notiophilus aesthuans (Mot- schulsky) Eee nno Notiophilus aquaticus (L.)..... Notiophilus biguttatus (F.).... Notiophilus germinyi Fauvel Notiophilus palustris (Duftschmid)................ Notiophilus rufipes Curtis..... Notiophilus substriatus Wa- TEFHOUSE AA n Odacantha melanura Paykull Olistophus rotundatus (Pay- Kai AA re: Omophron limbatum (F.)...... Oodes helopioides (F.).......... Ophonus azureus (F.)........... Ophonus cordatus (Duftsch- MAIALE van Shea Ophonus melleti Heer......... Ophonus nitidulus Stephens 304 So Hu 1 sa ni 1 pm 4 IN lo 2 1 sa 1 Ga 12 sm 2 sa 1 San? 2 SC 2 Ih al 2 2 Ga 2 3 N 2 [one sm pm pm sm 2 sc 4 pm sa 4 li 3 Sam 2 In 2 GR RA Bl Bl D3 D1 B2 EU RA Bl B2 Al EU Bl Bl B2 H2 D1 RA EU D2 D3 B2 EU EU Bl RA RA Al D3 EU H2 B2 Al EU EU Bl EU D3 Bl RA Al G4 G2 RA Bl D1 RA Species Ophonus puncticeps Stephens Ophonus puncticollis (Pay- Kun Ea Ophonus rufibarbis (F.)........ Ophonus rupicola Sturm....... Panagaeus bipustulatus (F.).... Panagaeus cruxmajor (L.)...... Parophonus maculicornis (Duftschmid)................ Patrobus atrorufus (Stroem) Pogonus chalceus (Marsham) Pogonus luridipennis (Ger- MALO na lier) ER e lo Pterostichus aterrimus (Herbst) RER, Pterostichus cristatus (Du- four RATORI, Pterostichus cupreus (L.)....... Pterostichus diligens (Sturm) Pterostichus gracilis (Dejean) Pterostichus lepidus (Leske) Pterostichus macer (Mar- Sham) ner N Pterostichus madidus (F.)...... Pterostichus melanarius (Il- gen) een hal) Mean nn Ran ee Pterostichus niger (Schaller) Pterostichus nigrita (Paykull) Pterostichus oblongopuncta- CUS) en Pterostichus quadrifoveolatus Metzler ees ar Pterostichus strenuus (Panzer) eN Pterostichus vernalis (Panzer) en eee Pterostichus versicolor (Sturm) SN Stenolophus mixtus (Herbst) Stenolophus skrimshiranus Stephens Stenolophus teutonus (Schrank) RR reer Stomis pumicatus (Panzer)... Syntomus foveatus (Fourcroy) Syntomus truncatellus (L.)..... Synuchus vivalis (Illiger)....... Tachys parvulus (Dejean)...... Trechoblemus micros (Herbst)! n Trechus obtusus Erichson...... Trechus quadristriatus (Schrank) I terrence Trechus rubens (F.)............. Trichocellus cognatus (Gyl- lenhaly Rai ov: eA rer ANR ee Trichotichnus nitens (Heer) NUM 69 So Hu GR TRD li 2 RA rc 2 DI 4 Gl sa 2 Bl G4 li 1 DI rc 4 F2 Is 5 H4 Is 5 H4 SC G4 pm 5 Al lo 3 RA sc H2 Al 3 RA pm Al RA I M2 MID? EU 4 Al EU EU D3 sm B2 EU EU sm EU H2 sc 5 RA 4 RA D2 EU sa 2 EU EU li 1 RA EU EU EU Is RA pm Al F2 lo 2 D3 No 090 0 oo 00 \O 00 00 00 LA \O KO vv N \O 00 Volume 134 1991 Tiydschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 AL ih LES 79 Pa EIER) YIN or Published by the Nederlandse Entomologische Vereniging Tijdschrift voor Entomologie A journal of systematic and evolutionary entomology since 1858 Scope The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long tradition in the publication of original papers on insect taxonomy and systematics. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeography, or ethology and ecology as far as meaningful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually, such papers will only be published when space allows. Editors E. J. van Nieukerken (elected 1986) and J. van Tol (1985) Co-editors A. W. M. Mol (1990) and R. T. A. Schouten (1990) Advisory board M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London). The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the ‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society), Amsterdam. Editorial address c/o National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands. Correspondence regarding membership of the society, subscriptions and possibilities for exchange of this journal should be addressed to: Nederlandse Entomologische Vereniging c/o Instituut voor Taxonomische Zoölogie Plantage Middenlaan 64 1018 DH Amsterdam The Netherlands Subscription price per volume Hfl. 300.- (postage included). Special rate for members of the society. Please enquire. Instructions to authors May be obtained from the editors. Graphic design Ontwerpers B.V., Aad Derwort, 's-Gravenhage Tijdschrift voor Entomologie Contents of Volume 134 Articles 145 155 177 278 31 35 183 193 Alders, K.: see Turin Bedos, A. & L. Deharveng Cephalachorutes gen. n., a new genus of tropical Neanuridae (Collembola). Bellamy, C. L. A review of the Philippine species of the Coroebus Gory & Laporte genus-group (Coleoptera, Buprestidae). Boer, P. |. den: see Turin Chen, P.: see Nieser Deharveng, L.: see Bedos Duffels, J. P. Revision of the genus Champaka (Homoptera, Cicadidae) from Borneo and Sulawesi. Essen, S. van: see Turin Gielis, C. Pterophorus Schaffer in the Ethiopian region (Lepidoptera: Pterophoridae: Pterophorinae): new species and checklist. Gielis, C. Corrigenda Heijerman, Th.: see Turin Hensen, R. V. Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae). Huemer, P. Caryocolum delphinatella (Constant) sp. rev., a senior synonym of C. fiorii (Klimesch) (Lepidoptera: Gelechiidae). Laane, W.: see Turin Lansbury, I. Cuticular blades and other structures of Diaprepocoris Kirkaldy and Stenocorixa Horvath (Heteroptera: Corixidae). Li, X. Z. & L. Y. Zheng New species of the genus Megacoelum Fieber from China (Miridae, Heteroptera). McAlpine, D. K. Relationships of the genus Heterocheila (Diptera: Sciomyzoidea) with description of a new family. 47 201 210 211 69 269 279 75 Nieser, N. & P. Chen Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau Buton (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), |. Nieukerken, E. J. van & R. Puplesis Taxonomy and distribution of the Trifurcula (Glaucolepis) raikhonae group (Lepi- doptera: Nepticulidae). Nieukerken, E. J. van Corrigenda Oosterbroek, P. & Br. Theowald Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera), with an index to the literature except Tipulidae. Penterman, E.: see Turin Puplesis, R.: see Nieukerken Puplesis, R., S. Seksjaeva & V. Sruoga Leaf-mining Lepidoptera (Nepticulidae, Bucculatricidae, Gracillariidae) from Ul- mus in northern Caspiya (Kaspia). Seksjaeva, S.: see Puplesis Sruoga, V.: see Puplesis Stonedahl, G. M. Review of the Oriental genus Angerianus Distant (Heteroptera: Miridae). Theowald, Br.: see Oosterbroek Turin, H., K. Alders, P. J. den Boer, S. van Essen, Th. Heijerman, W. Laane & E. Penterman Ecological characterization of carabid species (Coleoptera, Carabidae) in the Netherlands from thirty years of pitfall sampling. Vondel, B. J. van Revision of the palaearctic species of Haliplus subgenus Liaphlus Guignot (Coleop- tera: Haliplidae). Zheng, L. Y.: see Li Book reviews 68 74 Medvedev, G. S. (ed.), 1990. Keys to the Insects ofthe European Part ofthe USSR. Volume IV. Lepidoptera, Part 2. English Translation. [E. |. van Nieukerken]. Emmet, A. M., 1991. The scientific names of the British Lepidoptera. Their history and meaning. [E. |. van Nieukerken]. ® Rubtsov, |. A., 1990. Blackflies (Simuliidae). - Fauna of the USSR. Diptera, Vol. 6, Part 6. []. van Tol]. 154 Kime, R. D., 1990. A provisional atlas of European Myriapods. Part |. - Fauna Europaea Evertebrata, Volume |.[P.]. van Helsdingen). © Cherepanov, A. |, 1990. Cerambycidae of Northern Asia. Volume 1-3. [J. van Tol]. ® Veeresh, G. K., B. Mallik & C. A. Viraktamath (eds.), 1990. Social insects and the environment. Proceedings of the | Ith International Congress of IUSSI 1990. 200 Knight, W. J. & J. D. Holloway (Eds), 1990. Insects and the rain forests of South East Asia (Wallacea). [J. van Tol]. 268 Moller Pillot, H. K. M. & R. F. M. Buskens, 1990. De larven der Nederlandse Chironomidae (Diptera). Deel C: Autoekologie en verspreiding. [A. W. M. Mol]. Referees for volume 134 R. A. Angus (London), B. Aukema (Wageningen), M. Brancucci (Basel), G. W. Byers (Lawrence), W. N. Ellis (Amsterdam), |. K. Gelhaus (Philadelphia), P. |. van Helsdingen (Leiden), R. de Jong (Leiden), I. Lansbury (Oxford), M. L. Luff (Newcastle upon Tyne), A. Menke (Washington), N. Nieser (Tiel), R. Rozkosny (Brno). Dates of publication Volume 134 (1), pages 1-144, | July 1991 Volume 134 (2), pages 145-304, 18 December 1991 © Nederlandse Entomologische Vereniging, Amsterdam ISSN 0040-7496 NEW TAXA DESCRIBED IN TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134 COLEOPTERA Cisseicoraebus bicoloratus Bellamy ............ 163 Gissergagquilonta Bellamy nee 171 Coroebus albiventris Bellamy .................. 159 Coroebus bivestitus Bellamy .................... 159 Hahplussansusı vanıVondeln nenn. 85 Haliplus astrakhanus van Vondel ............... 85 Haliplus david: van Vondel ...................... 92 Haliplus excoffieri van Vondel .................. 96 Haliplus gafnyi van Vondel ............-........ 105 Haliplus holmeni van Vondel … … … … … … 109 Hahplusyaecht van Vondellerre een 109 HahplussortalisanıVondee nenne. 121 Hypocisseis philippinensis Bellamy ........... 174 BhiloconoebusBellamyame ee 164 Philocoroebus adamantinus Bellamy .......... 169 Philocoroebus alius Bellamy .................... 169 Philocoroebus elongatus Bellamy .............. 169 Philocoroebus maquilingensis Bellamy ....... 167 Philocoroebus pseudocisseis Bellamy ......... ILA Philocoroebus purpureus Bellamy ............. 168 Philocoroebus samarensis Bellamy ............ 167 Vanroonta luzonica Bellamy ...................- 175 COLLEMBOLA Cephalachorutes Bedos &Deharveng ......... 145 Cephalochorutes asiaticus BedosgegDehatvenee re E 152 Cephalochorutes barthae BedosaDehanven en 149 Cephalochorutes caecus Bedosgsa DEN AMEN NMR 149 Cephalochorutes centurionis BedosteaDehanveng rer RARE eue 149 Cephalochorutes murphyi BedosrsaDehanvemeune PEN RE ee 151 Cephalochorutes pestilentiae BeEdOS A DNA Ven. e 15 DIPTERA kleterocheilidae MeAlpiner rem 198 HETEROPTERA Angerianus mindanao Stonedahl .............. 275 Angerianus palawanensis Stonedahl .......... 275 Angerianus pallidus Stonedahl ................. 276 Aphelocheirus breviculus Nieser & Chen ...... 49 Aphelocheirus robustus Nieser & Chen ........ 50 Coptocatus stereos Nieser & Chen .............. DZ Enithares caesaries Nieser & Chen ............. DO, Enithares lansburyi Nieser & Chen ............. 62 Enithares phenakismos Nieser & Chen ........ 63 vi Enithares skutalis Nieser & Chen ............... 63 Megacoelum chinensis Li & Zheng ........... 185 Megacoelum pronotalis Li & Zheng .......... 184 Megacoelum pseudopronotalis Li & Zheng 187 Megacoelum rubripedum Li & Zheng ......... 185 Megacoelum tenwicorne Li & Zheng .......... 189 Megacoelum yunnananum Li & Zheng ....... 191 Megacoelum zoui Li & Zheng .................. 189 Ranatra sulawesti Nieser & Chen ............... 55 HYMENOPTERA Isodontia capillata Hensen ....................... 15 Isodontia cestra Hensen .......................... 15 Isodontia delicata Hensen ........................ 17 Isodontia elsei Hensen ............................ 19 Isodontia immaculata Hensen ................... 17 Isodontia nidulans Hensen ....................... 20 Isodontta Papua RSeRE er PEER ES 18 Isodontsarpilipes Hlensen en 13 SDNERTERNEIENSENS Ar EEEN 27 SIDES solommonALIEnSEenW 00000000090 25 SDhexawalshaelensen re 28 SDICIALSONMTE SEMPRE AIR DS) LEPIDOPTERA Bucculatrix caspica Puplesis & Sruoga .......... 71 Pterophorusdallastas Gielis 7 Pierophorusiuzunowe Giclissa eee eee 7 Piero phonasamassan CIC n 7 Stigmella kazakhstanica Puplesis ............... 70 Trifurcula (Glaucolepis) melanoptera van Nieukerkenié@ Puplesism- as ee 205 INSTRUCTIONS TO AUTHORS The Tijdschrift voor Entomologie publishes original papers dealing with systematic and evolu- tionary entomology. The editors particularly invite papers on the insect fauna of the Palaearctic and Indo-Australian regions, especially those including evolutionary aspects e.g. phylogeny and biogeo- graphy, or ethology and ecology as far as meaning- ful for insect taxonomy. Authors wishing to submit papers on disciplines related to taxonomy, e.g. des- criptive aspects of morphology, ethology, ecology and applied entomology, are requested to contact the editorial board before submitting. Usually such papers will only be accepted when space allows. Papers in English are preferred, but papers writ- ten in French or German will also be considered. It is our policy that papers are reviewed by an external referee. Authors will generally be notified of acceptance within two months. For the first submission two printed copies (or photocopies) are required, including photocopies of figures, reduced to A4 format. Diskette and orig- inal artwork should not be sent before the paper is accepted. Manuscripts should preferably be printed on A4 size paper, on one side only, double spaced, with a left margin of at least 4 cm. Authors not able to use a wordprocessor, are required to submit a neatly typed original, conforming to the same standards, without any handwriting on it; any re- marks should be written on a separate piece of paper. After acceptance of a paper, authors are requested to prepare a manuscript according to standards given in a separate style-sheet. These instructions include e.g. the wordprocessing pro- grams we can handle and the codes that should be included or are allowed in your files. The editors can handle the most common wordprocessing pro- grams for MS-DOS and Macintosh systems, but prefer Wordperfect (any version) or MS-Word. Lengthy papers (more than 40 pages in print) are only accepted when space allows. Publication can be speeded up by paying a page-charge of Hfl 50. Otherwise no page-charge is asked. Member- ship of the ‘Netherlands Entomological Society’ is not obligatory. Text preparation A cover page should provide the names of the authors and a proposal for a running title. The second page starts with author names (in all cap- itals), use & for ‘and’, on a new line the name of institute (as short as possible), with multiple au- thors using superscript ! ? etc. The title is brief and informative, typed in all capitals, with order and family of the taxon treated in parentheses. The abstract starts with a bibliographical refer- ence (Authors, year, title), see recent issues for layout. Then the abstract follows, without the word ‘abstract’ and without indentation. The abstract, written in english, should be concise, yet cover the main results of the paper, including new taxa and nomenclatorial changes. The name and address of one and only one of the authors follows, if needed preceded by the word ‘Correspondence:. This should also be the address for the galley proofs. The abstract ends with a list of keywords. The text proper starts on a new page, the intro- duction (if any) starts without heading. Use a max- imum of three categories of headings, all to be typed left, and using capitals and lower-case letters. The second and third type may be typed in bold, after the third type the paragraph follows without blank. A fourth category may be used in descrip- tions of species, etc. It follows an indentation and is separated from the following paragraph by a long dash (-), to be typed as double dash. All new paragraphs are indented: use a [TAB] code in wordprocessors. Scientific names of genera and species should be underlined. Although they will be printed in italics, no italics should be used in your wordprocessor. No underlining or italics are allowed for any other text. References In the text they are given as Lopes (1982a), (Lopes 1982) or (Brown & White 1975: 24). All cited papers should be listed alphabetically at the end of the paper under the heading ‘References’, papers not cited in the text should be omitted from the list of references. Examples for format: Boer, P. J. den, 1970. On the significance of dispersal power for populations of carabid-beetles (Coleoptera, Carabidae). - Oecologia 4: 1-28. Karsholt, O. & E. S. Nielsen, 1976. Systematisk forteg- nelse over Danmarks sommerfugle. - Scandinavian Science press, Klampenborg, 128 pp. Johansson, R. & E. S. Nielsen, 1990. Tribus Nepticulini. - In: Johansson, R. et al. The Nepticulidae and Opos- tegidae (Lepidoptera) of NW Europe. - Fauna ento- mologica scandinavica 23: 111-238, pls. Titles of journals should not be abbreviated. Type long dashes as double dash ‘- -’. Do not try to type indentation, just end each reference with a hard return. vii Nomenclature The latest edition of the ICZN Code should be followed. The composition of new names should preferably be explained in a paragraph ‘Etymol- ogy, including indication of gender of generic names and kind of specific name (adjective, noun in apposition, etc.). Use standard abbreviations : Sp. n., gen. n., comb. n., syn. n., sp. rev., nom. n., etc. For all genus and species-group names the authority (preferably with year of description) should be mentioned once. Author's names are not abbreviated. In new taxa the type material should be listed immediately after the name. Only holotype, lecto- type, neotype, paratype and paralectotype are al- lowed. Label data should not be quoted literally (except for primary types), but arranged in a stand- ardized sequence. Material should be listed alpha- betically under the present day countries or other geographical units. Long lists of non-type material should be summarized. Geographical names should be written according to present day spelling, orig- inal spelling or label names may be given in brackets. Use standard transcription for non-latin scripts (e.g. Pinyin for chinese, BSI for cyrillic, etc.) or refer to the ‘Times Atlas of the World’. Abbreviations (Codens) for depositories prefer- ably follow Arnett & Samuelson (1986: The insect and spider collections of the world. EJ. Brill/Flora & Fauna publications, Gainesville). Otherwise, they should be listed under ‘Material and methods’ or in the introduction. Data for primary types of previously described species follow directly the reference to the original description as: Elachista subnitidella Duponchel, [1843]: 326, pl. 77: 8. Lectotype & [designated by van Nieukerken & Johans- son 1987: 471]: [Austria, Vienna region], Duponchel coll., Genitalia slide EvN 2522 (MNHN) [examined]. Illustrations All illustrations, including photographs, graphs, maps, etc. should be serially numbered as figures. No subdivision with letters is recommended. Illus- Vill trations are to be reduced to column width (65 mm), 1.5 X column width (102.5 mm) or text width (135 mm). Line figures should be mounted in blocks, or are printed singly. When all figures are mounted in full-page blocks (after reduction: 135 X 195 mm including caption), they may be printed after the text, otherwise the approximate place in the text should be indicated with pencil in the margin of the manuscript. Line-drawings are num- bered with pre-stencilled or pre-printed figures, which should not be too large after reduction, pref- erably using a font like ‘Garamond’ or ‘Times’. Photographs should be unmounted glossy prints. Numbering of photos should be left to the discre- tion of the editors. Captions should be typed on a separate sheet (or in a separate file), consult latest issue for style. Colour plates will only be printed at the author’s expense. Tables Tables should be typed on separate sheets (or files), starting with the captions. When using a wordprocessor: start with a practical TAB setting, and use only one [TAB] code for each next column. No formatting with spaces is allowed. No lines should be added. Extensive and long tables should be avoided. Proofs, reprints Authors receive one proof only, which should be corrected and returned immediately. When correc- tions are few, sending per telefax is recommended. Authors receive 50 reprints free of charge. Ad- ditional reprints can be ordered when proofs are returned. Members of the Netherlands Entomo- logical Society receive a considerable discount. Cov- ers can be ordered at extra cost. All correspondence should be addressed to: Tijdschrift voor Entomologie, editors, attn. E. J. van Nieukerken / J. van Tol, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, Netherlands (Phone +31-71-143844, telefax +31- 71-133344). Tijdschrift voor Entomologie Volume 134, no. 2 Articles 145 155 177 183 193 201 210 211 269 278 279 A. Bedos & L. Deharveng Cephalachorutes gen. n., a new genus of tropical Neanuridae (Collembola). C. L. Bellamy A review of the Philippine species of the Coroebus Gory & Laporte genus-group (Coleoptera, Buprestidae). J. P. Duffels Revision of the genus Champaka (Homoptera, Cicadidae) from Borneo and Sulawesi. X. Z. Li & L. Y. Zheng New species of the genus Megacoelum Fieber from China (Miridae, Heteroptera). D. K. McAlpine Relationships of the genus Heterocheila (Diptera: Sciomyzoidea) with description of a new family. E. J. van Nieukerken & R. Puplesis Taxonomy and distribution of the Trifurcula (Glaucolepis) raikhonae group (Lepidoptera: Nepticulidae). E. J. van Nieukerken Corrigenda P. Oosterbroek & Br. Theowald Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera), with an index to the literature except Tipulidae. G. M. Stonedahl Review of the Oriental genus Angerianus Distant (Heteroptera: Miridae). C. Gielis Corrigenda H. Turin, K. Alders, P. J. den Boer, S. van Essen, Th. Heijerman, W. Laane & E. Penterman Ecological characterization of carabid species (Coleoptera, Carabidae) in the Netherlands from thirty years of pitfall sampling. Book announcements and reviews 154 200 Kime, R. D., 1990. A provisional atlas of European Myriapods. Part |. - Fauna Europaea Evertebrata, Volume |. [P. |. van Helsdingen]. © Cherepanov, A. I. 1990. Cerambycidae of Northern Asia. Volume |-3. []. van Tol]. ® Veeresh, G. K., B. Mallik & C. A. Viraktamath (eds.), 1990. Social insects and the environment. Proceedings of the | Ith International Congress of iussi 1990. Knight, W. |. & J. D. Holloway (Eds), 1990. Insects and the rain forests of South East Asia (Wallacea). []. van Tol]. Moller Pillot, H. K. M. & R. F. M. Buskens, 1990. De larven der Nederlandse Chironomidae (Diptera). Deel C: Autoekologie en verspreiding. [A. W. M. Mol]. Nederlandse Entomologische Vereniging, Amsterdam Published 18 December 1991 ISSN 0040-7496 pe gi 1: ra à RE PT RENE he 1 À Nr ii 1 LINE Y HERE 7 +t i , DI \ | 7 Bi) x Vi Al uk x ‘ NT 3 2044 093 383 164 eenma me ran meh gerne ae SES OU SE AN SP D ER NENTS OM ASTER TA rege tre ger Pag PB esta n na HEN impera VE Na REN remde ein tains E A ET TA MN È SP PS ore ee I Carni VARI ERTEILT wete ee arpa GE opsteeg N bressen re or EE MA eT NITY Ge RE RQ EN GO DeMille de onthaal ye WPAN tS dr rita ANE RISE ennn Zante bieden Vel PET urnen egt amen lenge gen DORE DRE VA erge AR SU UE ITR Er € = SEL A BRENG A TN Sa PEL TR eg mr LE Sie e Mea n n REA ot Oe avere en warn intra) AMIR FRI NAME ee AR wenende PEN eigen kn hee PSE Gre pigli astenga an Oe to ein dre Dar erster pre ua atri reni pretore TETTI NT RS PNR NG ONE eten tn RSR SR SU era DAS ES Beene meneer et ar eg men verteren vennen ra en Bare ar ee! mgee sen verten WETTEN UP 4 es “ur ke ATLANTE TA ALI earned A er AA oer auge hrs En RAN ER RUI LB Le PORT RHEE VERS Me VI ORI ente NS PANE ne WA NE PAN OOG MEIN USI ERN REEN era eR EN FRE AI MPI NERA D A TANT Ce Vire GH SATA RT Breeg Me TE ERS EPO RUE UL moh geisers er waren ree A Werner Apr ere UP Sebens serre ri Ur acter ea ek mye bieren Bevin ke! ET ACTEN A AIR OL es DDR peen netti RENTE DÉTENTE een Lee tn U ON AVE de METIN EN Gnd ener een Ly PN RLS A peen TS TANA A ENN SES EENES UL NET erp gate STA Ne EMSA ET RI HOE gd GENS maj re DA ba UIE Sa MEERLE Pa AU TE ENE Nete wf man pek PFA canes aI Den rein ra RIVE SE VA TIT en PT a VL ET Anker u RER RI VET PIPE re VA ADS DORE FRAZ TETI È ee TAN Braap dar warten ur GE as PATATE NEI da) BVLG ser PATES gui TP ed oF Hy Said elia RETE TE TORTI PETIT ARE san. ps DES aa gag dn dg iep nd oen agp ESSE IU pees pdre garde Bran a Ga OP TOAD RAI 5 è De Zar QUTE LIDL AR, ereen IE ne 0 Kd ta gemeen pane: erdee enge er raw wagons npe gegen M en vert