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A. W. Jeekel Vice-voorzitter (Vice-President) ......: NIRO L. H. M. Blommers Secretarisi(Secretamy) pren R. de Jong Address en CRT E, o re E Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden 2311 PL lerenninemeesten (easier Der ee L. P. S. van der Geest AGES STRA I MO Doornenburg 9, Landsmeer 1211 GP DelRenninemeestenidircasure dl) ARRESE P. Oosterbroek AGANESS RR RE SIR E N I Baanstraat 2, Edam 1135 CB Bibliothecanisi (librarian) ieee en ee ee W. N. Ellis Ae N AI Plantage Middenlaan 64, Amsterdam 1018 DH LENTE RO Ta B. van Aartsen TIJDSCHRIFT VOOR ENTOMOLOGIE RedactelEditonalBoand) ere me P.J. van Helsdingen, R. de Jong, J. Krikken, C. van Achterberg, S. A. Ulenberg, J. van Tol Addresse I tae ah sume li Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden 2311 PL The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. Subscription rate: D.Fl. 300,— per year. Issues 1—4 appeared on 20.X11.1985 ISSN 0040-7496 INHOUD Cobben, R. H. — Additions to the Eurasian saldid fauna, with a description of fourteen new spe- cIestl(hleteropteransaldid ao) Hr a An EU En 215 Jong, M. R. de. — Taxonomy and biogeography of Oriental Prasiini 1: the genus Prasia Stal, SOS (lomo preravmllibrermidac) Memmi samen ne EOS ai eA yh een 165 Nieukerken, E. J. van. — A taxonomic revision of the Western Palaearctic species of the subge- nera Zimmermannia Hering and Ectoedemia Busck s. str. (Lepidoptera, Nepticuli- dae)swithinotesionitheinphvlosenys FERIRE RE 1 Roskam, J. C. — Evolutionary patterns in gall midge — host plant associations (Diptera, Cecido- TAC OR E sik FACE AE FAO Me ENO AZ AES SR TRO CI nS ET 193 DEEL 128 AFLEVERING 1 1985 TIJDSCHRIFT VOOR ENTOMOLOGIE “UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING f 101 6 \ 3 | iw Si i | 1200 F \ J INHOUD E. J. van NIEUKERKEN. — A taxonomic revision of the Western Palaearctic species of the subgenera Zimmermannia Hering and Ectoedemia Busck s.str. (Lepido- ptera, Nepticulidae), with notes on their phylogeny, pp. 1—164, figs. 1—549. Tijdschrift voor Entomologie, deel 128, afl. 1 Gepubliceerd 20-XII-1985 eb tah a ad PE NI x Ret DROPS BN dites Line. ah, ANS s A TAXONOMIC REVISION OF THE WESTERN PALAEARCTIC SPECIES OF THE SUBGENERA ZIMMERMANNIA HERING AND ECTOEDEMIA BUSCK S.STR. (LEPIDOPTERA, NEPTICULIDAE), WITH NOTES ON THEIR PHYLOGENY by ERIK J. VAN NIEUKERKEN Department of Animal Systematics and Zoogeography, Vrye Universiteit, Amsterdam, Netherlands ABSTRACT The subgenera Zimmermannia Hering and Ectoedemia s.str., together forming the genus Ectoedemia Busck sensu Wilkinson & Newton (1981) are described and redefined, and the Western Palaearctic species are revised. In total 50 species are recognised, including the new species hispanica, monemvasiae, nuristanica in Zimmermannia and andalusiae, algeriensis, leucothorax, alnifoliae, contorta and two unnamed species in Ectoedemia s.str. Fifteen new synonymies and ten new combinations are established and 42 lectotypes are designated. Primary types have been examined in many cases. Data on larvae and biology are included and keys to all species are provided. The monophyly and the sister group relationships of both subgenera are demonstrated. The subgenus Ectoedemia can be divided into the populella group, suberis group, subbima- culella group and occultella group, being monophyletic entities, and the possibly paraphyle- tic angulifasciella group. Two alternative hypotheses of the phylogeny within Ectoedemia s.str. are presented. Decisions on species discrimination have in many cases been corroborated by study of allozymes. CONTENTS Weknowledvements ssa ae er ee eee re 92 RDS BAR ee Si ue te a Me aie 1", References JMeter hia Oi A ers I 92 AO ETD ree ee ee ee 1 Index to (sub)genera and species treated ........ 98 REESE BEE RIO AERO CARE CES 4 Robart polo payer E gl ren RENE GR re ee : INTRODUCTION Hiaxonomic treatments: us). bi Sash ee ee 8 The present revision deals with the 50 West- Checklist of species treated .. SOCCER eee eee 8 ern Palaearctic species of Ectoedemia Busck, Keys to the Western Palaearctic species of Ectoe- 1907, here assigned to the subgenera Zimmer- or ame Zimmermannia and Ectoe- 9 mannia Hering and Ectoedemia s.str. These two Subgenus Zimmermannia .................... 17 form the genus Ectoedemia in the sense of Wil- LENS ÉOLIEN 27 Kinson & Scoble (1979) and Wilkinson & New- Whepopwlellgeroupls en, A NO UNE SORA 28 ton (1981). The concept of Ectoedemia was re- Thepreisseckerügroup . NON 37 cently enlarged by Scoble (1983) to contain the Whetwberislevoupss gern GAL ai 38 subgenera Fomoria Beirne and Laqueus Scoble, The subbimaculella PROUP TAS MERE RIAA o 43 and one more subgenus will be included in a Theterebinthivora group … sooner 63 forthcoming generic revision of Holarctic Nep- Dal angalizsciella CLOUD Me EE REC CI erlas (Van Nieukerken, in preparation). An NG CCOAMALR LOUD Se EEE ES Come 78 lee EN Pal He Names of doubtful status, probably belonging to oh Bo ER are De Era EE g2 Species assigned to the subgenera of Ectoedemia Catalogue of Hostplants of Western Palaearctic ger treated here, will be presented by Van ILL OE Dd BIS EAD ie ee EL Baa ON 82 Nieukerken (in press). PRIS STORE DR Eee BILIA ea 85 Throughout this work the name Ectoedemia Bioseosraphyan re RIN a ee 91 alone is reserved for the combination of the two 2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 subgenera treated here, Ectoedemia s.str. is the typical subgenus and Ectoedemia s.l. is the en- larged genus in the concept of Van Nieukerken (in press). For a taxonomic history of the genus and de- scription of the Nearctic species refer to Wil- kinson & Scoble (1979) and Wilkinson & New- ton (1981). The three South-African species have been described by Scoble (1978, 1979). A complete revision of the known Ectoede- mia species in the Western Palaearctic region has not been carried out previously. The last au- thor reviewing all European species was Meess (1910) who assigned most species to Nepticula Heyden (= Stigmella Schrank). Of the 138 spe- cies in his work only 19 belong to Ectoedemia in the present sense, of which 15 are here recog- nised as good species, hence the number of spe- cies has since been more than tripled. In Europe Petersen (1930) figured the male genitalia of some nepticulid species for the first time, but retained them in the large genus Nep- ticula. Beirne (1945), who divided the Nepticu- lidae into several genera on the basis of the male genitalia of the British species, erected the genus Dechtiria for the leafmining species here as- signed to Ectoedemia s.str. Hering (1940) erected Zimmermannia as a genus for the barkminers, but most European authors placed them in Ectoedemia, following Busck, as did Klimesch (1953) in his revision of the four known European species. Svensson (1966) was the first to discover the similarity between Ectoedemia and Dechtiria and hence synonymised both. This was fol- lowed by Borkowski (1972) and Emmet (1976) in their local fauna works. These authors recog- nised Dechtiria and Zimmermannia as separate subgenera, but Wilkinson & Newton (1981) treated them as synonyms of Ectoedemia. Zim- mermannia is here re-established as subgenus for reasons to be discussed. Apart from the four species treated by Kli- mesch (1953), no part of the genus has been completely revised and published in Europe previously. Most species described since Meess (1910) were assigned originally to Nepticula or Stig- mella, but several have in recent years been re- combined with Ectoedemia or Trifurcula s.l., al- though frequently only in faunistic lists, with- out any comments. Most names given to European nepticulid species have been assigned to their correct genus in my checklist (Van Nieukerken, in press), but a few doubtful names still exist. Two are treated at the end of this pa- per. For those species, likely to be included in Ect- oedemia, primary types were studied as far as possible. A few old types were either not avail- able during this study or could not be traced. In most cases however, there has been enough proof of their status. For some recently de- scribed species, no types have been studied, be- cause detailed description and figures of genita- lia made it unnecessary. A large wealth of material from several museums, private collec- tions and our own collection has been studied and resulted in the discovery of eight undes- cribed species, and much new distribution data. However, knowledge of Ectoedemia species in the Mediterranean region and Middle East is still poor and based on scanty data, as can be in- ferred from the distribution maps. For instance none of the autumn-feeding species of the an- gulifasciella group are recorded from Spain, probably because autumn-mines have not yet been collected. For all species, including those recently de- scribed, complete (re)descriptions are provided. For most species the female genitalia are de- scribed here for the first time. These often ap- pear to give better diagnostic characters than the male genitalia in this genus. Because of limitations in time and space, I have refrained from giving detailed descriptions of larvae, although much material was available. However, it is hoped that a full treatment of the larvae can be made later. _ Concise biological data have been provided, based on own observations, unless otherwise stated. A discussion of the phylogeny of the genus, using cladistic methods, concludes this revision. METHODS Preparation of genitalia Genitalia slides were prepared following Robinson (1976), but adapted slightly for the Nepticulidae. The abdomens were macerated in 10% KOH heated in a waterbath of 90 °C for 10—15 minutes. After preliminary rinsing and cleaning they were stored overnight in ethanol 70%. Cleaning appeared to be much easier after treatment with ethanol and there were no disad- vantages. Cleaning and removal of scales was carried out with a snipe-feather primary or a pointed piece of stiff paper. For dissecting min- ute-pins were mounted in handles. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 3 Male genitalia were usually stained red with haemaluin and females either with haemaluin or chlorazol black E. Dissecting was usually done in glycerin to prevent floating. Male genitalia were removed from the abdomen, the aedeagus was taken out in some specimens of each species, by perforat- ing the membranes holding it to the valvae and capsule; in Ectoedemia this is often difficult be- cause of the tight connections to the aedeagal carinae. It is therefore advisable not to remove the aedeagus from all specimens, otherwise their in-situ connections can not be studied. Hooking out the vesica is possible in the larger species, but usually impracticable in smaller ones. Fe- male genitalia were removed by separating seg- ment 7 to 9 with the internal genitalia from the abdomen. Before mounting, genitalia were ex- amined in glycerin in order to study their three- dimensional structure and to make figures in various aspects. After dehydration the genitalia were em- bedded in euparal, and arranged in their desired position. The euparal was placed in a thin layer so that the parts could not move and the slide was then dried in an oven overnight. Thereafter a small drop of euparal was added and the cov- erslip positioned with, if necessary, euparal es- sence. This method prevents the parts from be- coming displaced and disorientated. Care must however be taken not to damage protruding parts such as the gnathos or uncus with the cov- erslip. Male genitalia were mounted ventral side up, female genitalia either with ventral or dorsal side. In order to study the female postabdomen embedding with dorsal side up is most desir- able. In the above described method the genitalia are not squashed, which has a disadvantage in that focussing for photography is difficult, but this is outweighed by the disadvantage of dis- tortion by squashing. It has unfortunately proved to be virtually impossible to unroll the male genitalia in the way practiced for Incurva- rioidea (see Nielsen, 1980), because of the strongly sclerotised capsule, the tightly fused valvae, and the small size of the genitalia. Figures Drawings of genitalia were made with a Zeiss universal microscope and camera lucida attach- ment both from permanent slides and genitalıa in glycerin. Dorsal aspects of valvae were drawn from ventrally mounted specimens, thus repre- senting in fact a mirror image of the right valva as seen through the valva. From the transtilla only one half is figured. Setae are often repre- sented in drawings by their sockets only be- cause they are often broken in slides. In the fig- ures of aedeagi in Ectoedemia s.str. the vesica is omitted. The practice of illustrating complete genitalia in taxonomic papers on Lepidoptera is not fol- lowed, since such figures are usually too com- plicated to show the diagnostic features unam- biguously. Therefore the most characteristic parts of the genitalia are separately figured and presented in a comparative way. However, to give an overall impression of the genitalia, pho- tographs are also provided. These were pre- pared with a Zeiss universal photo-microscope, using bright-field contrast. SEM micrographs were taken with an ISI 40 Scanning electron microscope, using a beam current of 10kV. Specimens were air-dried, mounted on stubs and gold-coated. Adults were photographed with a Zeiss Tes- sovar camera, using black velvet as background, and concealed lighting, thus reducing reflections to a minimum. Photographs of mines in dried leaves were taken with a reproduction camera and transmitted light. Measurements Forewing length was measured only when flat from wing base to tip of fringe, using an oc- ular-micrometer in a Wild M5 stereomicroscope at a magnification of 25. Forewing length is pre- ferred to the less accurate wingspan mea- surement, but for reasons of comparability with other authors the latter figure is added too. Genitalia were measured using a Zeiss univer- sal research microscope with ocular-microme- ter, either with objective 6.3 X (bursa length and signa if very long) or 16 X (other mea- surements). Capsule length was measured along mid-line from tip of tegumen to anterior margin of ventral plate of vinculum, exactly in middle of anterior concavity, thus excluding lateral projections of vinculum. Valva length was mea- sured from tip to anteriormost extension of ven- tral surface, thus excluding the transulla. Ae- deagus length was measured including carinal processes. The bursa length could only be measured very roughly, approximately from point of en- trance of ductus spermathecae to anterior tp. Measurements of signa are self-evident. 4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 From all species measurement range is given first, followed by mean, standard deviation and sample-size in brackets. Mean and standard deviations are only calcu- lated for a sample-size of five and larger. An in- dividual of extreme size falling far outside the normal range is given in brackets. Wing mea- surements of extremely small specimens, proba- bly caused by food-shortage, are excluded. Not too much statistical significance should be given to these figures, because the samples were not selected statistically, and sometimes individuals only belong to one population. Material A considerable part of the adult material was reared in our laboratory, and will be mainly transferred to the collection of ZMA, however some specimens will be distributed to other mu- seums. In addition material of many collections, listed below, has been examined. The material is listed at the end of each description in alphabeti- cal order of localities, arranged in an alphabeti- cal list of countries. When a number of consec- utive data in one country is based on material from one collection, the abbreviation of this collection is only given at the end of these data. Primary types, cited under the species headings are included in material examined again, when actually studied. Locality names are spelled as far as possible according to The Times Atlas of the World (Comprehensive Edition, 1975), a deviating name on a label is given in brackets. A particular problem form the locality-names on the labels of C. Chrétien, who often used ab- breviations of small hamlets or local names, which can not even be traced on topographical maps. By courtesy of G. Luquet, who prepared a list of Départements visited by Chrétien in various years, it has been possible to locate some of these obscure places. «Antarv.» has not been traced, but the collecting dates suggest that this is near Digne. «Nesp.» is an abbreviation of Nespouls, but most likely not of the village of that name in Corréze. From a combination with “Artén.” (= montagne d’Arténac) and the col- lecting year on certain labels it is inferred to be probably near St. Pons (Hérault). Countries are used with their present-day po- litical boundaries, but for convenience East- Germany comprises here both the German Democratic Republic and Berlin. Distribution maps are prepared on the base of material examined and reliable literature re- cords. When a certain literature record was far beyond the known range, and its correctness could not otherwise be proved it has been ex- cluded. Many additonal data were received by courtesy of R. Buvat, R. Johansson, O. Kars- holt, J. Klimesch, J. Kyrki and S. E. White- bread. A list of literature used in compiling the maps wil be given later. The data on biology are for a considerable part based on own observations, supplemented by literature data. Unless otherwise stated, mines have been collected between 1978 and 1984 by me or my colleagues or students, chief- ly C.J. M. Alders, J. J. Boomsma, G. Bryan, B. J. van Cronenburg, H. van Driel, S. B. J. Men- ken, J. W. Schoorl, and stored in our collection. Larvae have been examined living, and are part- ly also stored in alcohol in our collection. Nomenclature of hostplants follows Tutin et al. (1964, 1968). Some abbreviations used are: © a.l. = at light, e.l. = ex larva, S = sternite, T = tergite. List of collections from which material has been studied Institutions and Museums: BMNH, British Museum (Natural History), London, U.K.; ETHZ, Eidgenössische Technische Hochschule, Entomologisches Institut, Zurich, Switzerland; IPAK, Institute of the Polish Academy of Sci- ences, Krakow, Poland; IRSN, Institute Royal des Sciences naturelles, Bruxelles, Belgium; LNK, Landessammlungen für Naturkunde, Karlsruhe, West Germany; MCST, Museo Civ- ico di Storia Naturale, Terrasini, Italy; MHUB, Museum ftir Naturkunde der Humboldt-Uni- versitat, Berlin, East Germany; MNHN, Muséum national d’Histoire naturelle, Paris, France; MRST, Museo Regionale di Scienze Naturali, Torino, Italy; NMW, Naturhisto- risches Museum, Wien, Austria; RMNH, Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands; RMS, Riksmuseum Stockholm, Sweden; SMNS, Staatliches Museum fur Natur- kunde, Stuttgart, West Germany; TMAB, Természettudomanyi Múzeum, Allatära, Buda- pest, Hungary; UMZC, University Museum of Zoology, Cambridge, U.K.; USNM, United States Natural History Museum, Smithsonian Institution, Washington D.C., U.S.A.; ZIAS, Zoological Institute, Academy of Sciences, Le- ningrad, USSR; ZMA, Instituut voor Taxono- mische Zoologie (Zoologisch Museum), Amsterdam, Netherlands; ZMC, Zoologisk Museum, Universitet, Kobenhavn, Denmark; Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 5 ZSM, Zoologische Staatssammlung, München, West Germany; ZSMK, idem, collection Kli- mesch, Linz, Austria. Private collections: AFW, coll. Van Franken- huyzen, Wageningen, Netherlands; coll. Buvat, Marseille, France; coll. Derra, Bamberg, West Germany; ETO, coll. Traugott-Olsen, Marbel- la, Spain; EvN, coll. Van Nieukerken, Leiden, Netherlands; coll. Gielis, Lexmond, Nether- lands; coll. Huisman, Melissant, Netherlands; coll. Johansson, Växjö, Sweden; coll. Koster, Callantsoog, Netherlands; coll. Kuchlein, Wa- geningen, Netherlands; coll. Leraut, Paris, France; coll. Speidel, Karlsruhe, West Ger- many; coll. Wolschrijn, Apeldoorn, Nether- lands. MORPHOLOGY The following discussion is mainly intended to review those characteristics which are impor- tant for understanding the phylogeny of Ectoe- demia and those which are useful as diagnostic features. Exhaustive treatments of the adult morphology of Ectoedemia and the Nepticuli- dae are given by Scoble (1979 and 1983), and of the larval morphology by Gustafsson (1981a) and Van Nieukerken & Jansen (in preparation). Schönherr (1958) provides an excellent mono- graph of the species E. liebwerdella. Head (fig. 15). The piliform scales on frons and vertex are collectively treated as the frontal tuft, the colour of which is often diagnostic, although some lo- cal and geographical variation occurs in several species. In Ectoedemia the collar is invariably composed of piliform scales, in contrast to Stig- mella where the scales are lamellar. Its colour 1s often different from the frontal tuft. The term collar, although descriptive, might be mislead- ing, since these groups of scales, inserted poste- rior of the eyes, are not homologous with the collar of higher Ditrysia, which is a prothoracic structure. The number of antennal segments has some diagnostic value, although it varies within a spe- cies and sex, males have always more segments than females of the same species. There is also a positive correlation between individual size and number of antennal segments. Scape and pedicel are usually paler than the flagel, except in E. ın- timella. For a detailed description of antennal morphology see Van Nieukerken & Dop (in preparation). The mouthparts of the species treated do not show diagnostic features. The eyes show the typical lepidopteran corneal nipple array pat- tern (fig. 16) (Davis, 1978). Thorax and wings. The thorax itself does not present many char- acteristics, the colour of the scales on mesoscu- tum and tegulae is sometimes diagnostic, but in many species it is concolorous with the fore- wings. The colour-pattern and colour of the fore- wings is one of the most remarkable diagnostic features, although it is only useful in undam- aged specimens, and many closely related spe- cies have the same or a similar colour-pattern. Most species of Ectoedemia s.str. have white wing markings, often in the form of a medial fascia, or opposite costal and dorsal spots. In addition basal and discal spots may occur. It is often difficult to distinguish between metallic and non-metallic fasciae and spots. Compari- sons should therefore be made with species in which this state is known. Several species, espe- cially in Zimmermannia, have the forewings uniformly ochreous irrorate with brown, fus- cous or similar tinges. In all but a few species the cilia are light and separated from the darker part of the forewing by a line formed by the tips of the last row of lamellar scales, this line is termed here the cilia-line. The scaling of the forewing is invariably rough, the scales (figs. 25, 26) are of the normal advanced lepidopteran type (Kristensen, 1970; Davis, 1978). The hindwing of the males frequently pos- sesses diagnostic secondary sexual characters. A frenulum is always present, in additon several species mining Quercus have a row of costal bristles. Most other species however bear a brush of hair-scales instead, arising near the fre- nulum, which is believed to be homologous with the costal bristles. Following Scoble (1983) it is named here hair-pencil. In rest it is laid par- allel to the main-axis of the hindwing, in a shal- low groove, which is especially prominent in several E. (Zimmermannia) species (figs. 10— 14, 21—24). The hair-pencil can be spread out, and probably plays an important role in courtship, as Schönherr (1958) has shown for E. liebwerdella (see his figs. 26 and 41). The hair- pencil is often surrounded by lamellar scales which are differently coloured from the rest of the hindwing, these scales are referred to as spe- cial or androconial scales (figs. 18—20). The fine structure differs from the normal wing scales. In some species they occupy almost the 6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 complete dorsal surface of the hindwing, as in terebinthivora or heringella (figs. 53, 62). Col- our of these scales and the hair-pencil is very di- agnostic. In E. (Zimmermannia) and to a lesser extent in some other species, the shape of the hindwing is influenced by the presence of the hair-pencil: the costal margin is abruptedly emarginated and curved inwards beyond the pencil and there is often a prominent humeral lobe (figs. 8, 10—19). In several species the males possess in addi- tion to the hindwing characteristics, specialisa- tions on the underside of the forewing, such as a patch of differently coloured, androconial scales (figs. 63, 86). Species with a hair-pencil often have a scaleless area on the forewing under sur- face, probably in rest contacting the hair-pencil. Females always bear a row of costal bristles, and lack any additional sexual characters. The venation of Ectoedemia (figs. 8, 9) is very uniform, with only slight non-diagnostic varia- tion in length and tracheation of some veins. The venation is essentially similar to that in the taxa Fomoria Beirne and Etainia Beirne. Abdomen. The scaling of the abdomen is uniform, and although there is some interspecific variation, the colour has not been found to be diagnostic. The anterior part of sternite 2 (see Kristensen & Nielsen, 1980) has a triangular shape. The male bears a pair of anal tufts on tergite 8. The exter- nal shape of the female ovipositor is sometimes diagnostic, especially when it is pointed, such as in E. turbidella or agrimoniae. Male genitalia (figs. 3, 4, 27, 28). The male genitalia of the species under study show a remarkable uniformity when compared to other nepticulid genera, in several cases they do not even provide characters to distinguish between species. It must be stressed here that slight differences which apear from the illustrations often depend on the way of mounting the slides. A slight de- viation from the ventral view can change the shape of the vinculum for instance, and since most structures are hinged by membranes to each other, mutual changes in position occur easily. This is especially the case with the gna- thos. It is therefore advisable to study the geni- talia in fluid (glycerin) before mounting perma- nently, and squashing should be avoided. The vinculum forms a complete strongly scle- roused ring and is invisibly fused with the tegu- men; together they are termed the capsule. The ventral plate of the vinculum is always short, and slightly concave anteriorly; the ventral plate can be divided by the ring, formed by the at- tachment to segment 8, in an anterior part, which is situated within the abdomen, and a posterior part, covered with scales. The anterior part has in the past erroneously been referred to as the saccus (Beirne, 1945; Wilkinson & New- ton, 1981). The tegumen is posteriorly produced into a pseuduncus, which can be approximately trian- gular, rounded, truncate or pointed. It is cov- ered with many tactile hairs and scales. The uncus is absent, it has been believed (cf Beirne, 1945) that it is membranous, but the membranous structure which is present be- tween gnathos and tegumen is in my opinion formed by the anal tube only. The gnathos is strongly sclerotised, and es- sentially composed of two lateral arms and a more ventral central element, which projects posteriorly and is more or less tongue-shaped. The form of the central element is highly diag- nostic, but it must be viewed at the correct an- gle. In several species of Ectoedemia s.str. the central element is in fact divided in two parts: a basal ventral part, fused to the lateral arms and distally ending with a serrate margin, and a more distal, tongue-shaped element which is in- serted dorsal to the basal part and connected by less sclerotised tissue. In lateral view the divi- sion is clearly seen, but in ventral view this is less obvious. The lateral arms of the gnathos are hinged by membranes to the lateral arms of the vinculum. The valva is roughly triangular in ventral view, with an often inwardly directed tip. It is essentially a hollow sac, which is open at the an- terior end. On the ventral and outer (lateral) surface, the valva is covered with many setae and scales, whilst the inner and dorsal surfaces bear comparatively few setae, which however become more abundant towards the tip. Al- though it has been the practice in Nepticulidae to illustrate only the ventral surface of the valva, the dorsal surface offers more diagnostic detail and so is here illustrated as seen through ven- trally mounted genitalia — thus viewing through the valva. Therefore it has not been necessary to spread or remove the valva. Ven- trally the valvae are hinged to each other and the vinculum by membranes, dorsally they are tightly fused by the transtillae, which are con- sidered to be a part of the valvae. In Ectoedemia ns Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 7 the transtillae always possess a well sclerotised horizontal bar and ventral arms. The length of the ventral arms varies within the species and therefore has a limited diagnostic value. The aedeagus bears apically paired carinae, except in E. spiraeae. These have been incor- rectly referred to as the juxta by Beirne (1945), see also Scoble (1983) for a discussion of aedea- gal structures. Most Ectoedemia s.str. species have one pair of ventral carinae only, they are usually pointed and often divided in two or more processes. Additional spines occur in some species near the base of the carinae (fig. 28). Some species have a dorsolateral pair of ca- rinae in addition, and most E. (Zimmermannia) species possess three pairs of carinae. The ven- tral carinae are hinged by a slightly sclerotised ventral process to the vinculum and by mem- branes to the valvae. Although the ventral pro- cess is always present, it has not been illustrated here in all species. Dorsal and lateral carinae are hinged by membranes to the valvae and transtil- la. In E. (Zimmermannia) the large ventral cari- nae are tightly connected with the valvae, which have a fold on the inner surface in which the ca- rinae fit. Probably for this reason the carinae in this group have sometimes been misinterpreted as parts of the valva. The membranes fusing the aedeagus to the rest of the genitalia tolerate only a small posterior movement of the aedeagus, hence in the everted position the carinae are folded back. The aedeagus is often slightly asymmetric, such that it is longer at the right side, and often the two carinae of one pair differ slightly. An exceptional case is klimeschi, which has a highly asymmetrical aedeagus. The ejaculatory duct enters the aedeagus through an approximately circular opening on the ventral side, below the middle. Posterior to this opening a group of mi- crosetae (setal pores) can be observed. The vesi- ca is typically covered with numerous small spine-like cornuti, and only occasionally addi- tional larger spines and other sclerotisations oc- cur. Female genitalia (figs. 6, 7, 30—34). The female genitalia of Nepticulidae have been paid much less attention to than those of the male, since they were often thought of lesser diagnostic value. In fact in Ectoedemia they of- ten provide better characteristics than the male genitalia. However, the weak sclerotisation and the greater individual variation — compared with males — make study and interpretation more difficult. Several structures, which have not been used before, are found in this study to have high diagnostic value. The only earlier complete and correct inter- pretation of the terminal segments is that of Dugdale (1974), see also the comments in Van Nieukerken (1983). Segment 7 is the last com- plete and more or less unmodified segment, which ventrally reaches the tip of the abdomen. The tip of sternite 7 is covered with many setae, probably mostly tactile. Dorsally tergite 7 en- circles segments 8 and 9. Segment 8 comprises a distinct tergite, which is often approximately rectangular, and the complex anterior apo- physes. They are dorsally united by tergite 8 and posteriorly by a semi-circular or angular, sclerotised bar, which is interpreted as sternite 8. The latter is covered by a membrane bearing many minute spines (fig. 30) and forms usually the tip of the abdomen and “ovipositor”. It is not completely clear if the integument covering tergite 8 belongs to that segment or is formed by segment 7, the latter possibility 1s suggested by the fact that the border between tergites 7 and 8 is often not clear. For practical reasons, however, setae and scales which in dor- sal view appear to occur on tergite 8 are de- scribed as belonging to that segment. The poste- rior part of tergites 7 and 8 bear several sensory structures. Principally there are two lateral patches of scales and setae on tergite 8, and of- ten some setae on tergite 7 as well, which is fur- ther covered with scales. In several species the scales on 8 are reduced and the number and size of setae increased, often forming distinct pat- terns or rows. Especially in species mining bark and evergreen Quercus there are large groups of long setae on these segments (figs. 31—34), which probably function in localising suitable oviposition sites. It is not clear if these setae are all mechanoreceptors only, or if these are partly chemoreceptors as well. In E. caradjai and E. monemvasiae the long setae are pectinate (fig. 32), in other species examined they are smooth. Segment 9 comprises a distinct tergite, often partly covered by tergite 8, with two distinct patches of setae (anal papillae) and the posterior apophyses. These end in indistinctly sclerotised internal structures, which probably have a func- tion in opening and closing the genital and anal openings. The region near tergite 9 is difficult to interpret since many membranous structures occur, it is therefore not clear if there is under- neath the anal opening a structure which can be considered to be sternite 9. 8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 The enlarged portion of the vagina is referred to as the vestibulum, this part has earlier been regarded as part of the ductus bursae (Scoble, 1983) and sometimes termed colliculum (Wil- kinson & Scoble, 1979; Wilkinson & Newton, 1981). Here the term ductus bursae is reserved for the narrowed part anterior of the entrance of the ductus spermathecae. The vestibulum bears in E. (Zimmermannia) some indistinct sclerotisations and in most Ec- toedemia s.str. species a ring-shaped sclerite (fig. 419), which in analogy to the Eriocraniidae (Davis, 1978) is termed vaginal sclerite. In addi- tion to this sclerite the vestibulum has dorsally an evaginated pouch which often bears many spicules (fig. 420). At the transition of the vesti- bulum and the ductus bursae there is often a patch of very closely packed pectinations, simi- lar to those on the corpus bursae. The corpus bursae is typically covered with many of such pectinations, combs of small denticles, and a pair of reticulate signa. The cells of the signa are also covered by small denticles. Shape and size of the signa is often diagnostic, but there is con- siderable intraspecific variation. The ductus spermathecae comprises a strongly sclerotised internal canal, ending in a sclerotised vesicle, and a membranous external canal, both canals are spiraled. The number of convolutions of the spermathecal duct appears to be fairly constant within a species, and has therefore high diag- nostic value. The most common condition is 2%—3 convolutions, but as many as 14 convo- lutions have been found. In counting the convo- lutions the vesicle must be excluded. A distinct spermathecal papilla is absent. TAXONOMIC TREATMENT Ectoedemia Busck, 1907 (subgenera Zimmermannia Hering and Ectoedemia s.str.) Diagnosis. The following combination of characters is diagnostic: 1. Collar comprising piliform scales. 2. Cilia-line usually distinct (except occultella- group). 3. Forewing with closed cel between R and M+Cu. 4. Hindwing with two-branched Rs+M. 5. Antenna with sensillum vesiculocladum re- duced into an unbranched blisterlike struc- ture (Van Nieukerken & Dop, in prepara- tion). 6. In © only 1 sensillum vesiculocladum per segment (Van Nieukerken & Dop, in prepa- ration). 7. Uncus absent. Stigmella species are easily separated by the collar with lamellar scales and the different ve- nation and genitalia. Acalyptris (= Niepeltia) species can be separated by the almost straight R+M vein in the forewing, and the reduced closed cell, shifted towards the base. Externally Acalyptris species in Europe are not likely to be confused with Ectoedemia because they have different colour patterns. Only A. minimella (Rebel) resembles somewhat E. gilvipennella or E. nigrosparsella, but it is more yellow and has a yellow hair-pencil. Trifurcula species can al- ways be recognised by the three branched con- dition of the Rs+M in the hindwing. In addition males can always be recognized by the three pairs of anal tufts and the “velvet patch” on the underside of the hindwing. For Parafomoria see Van Nieukerken (1983). European Ectoedemia (Etainia) species have two fasciae, and the males possess a long dorsal apodeme on the valvae (see Scoble, 1983). Bohemannia species can be sepa- rated by the absence of a closed cell in forewing and the presence of an uncus. Ectoedemia (Fo- moria) and E. (Laqueus) are externally not sepa- rable from the subgenera treated here. They both possess an uncus, and-have generally a dif- ferent form of genitalia (Scoble, 1983). In addi- tion E. (Laqueus) has an anal loop in the fore- wing. Taxonomy. Two subgenera are recognised here, viz. Zim- mermannia Hering and Ectoedemia s.str. This division is re-established here, because both groups are characterised by many more apo- morphies than they share, they have very differ- ent biologies, and species can easily be recogni- sed as belonging to one of the subgenera. Ectoe- demia s.str. can also be subdivided further, but then much fewer characters are available and monophyly is not easily demonstrated. These groups are merely treated as species-groups without formal taxonomic status. See further section on phylogeny. CHECKLIST OF SPECIES TREATED Ectoedemia Busck Subgenus Zimmermannia Hering 1. atrifrontella (Stainton) 2. hebwerdella Zimmermann 3. longicaudella Klimesch peinu (Nemes) syn. n. OND LA Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 9 hispanica sp. n. monemvasiae sp. n. amani Svensson nuristanica Sp. n. liguricella Klimesch Subgenus Ectoedemia Busck Dechtiria Beirne mes group 4 I. 12. 13. intimella (Zeller) hannoverella (Glitz) turbidella (Zeller) populialbae (Hering) klimeschi (Skala) niculescui (Nemes) syn. n. argyropeza (Zeller) simplicella (Heinemann) syn. n. preisseckeri group 14. preisseckeri (Klimesch) suberis group ley 16. 17. 18. 19. caradjai (Groschke) spec. (specimen 1843) suberis (Stainton) comb. n. viridella (Mendes) syn. n. andalusiae sp. n. aegilopidella (Klimesch) comb. n. subbimaculella group 20. 21: 22. DSE 24. 25: 26. 27: 28. 29. 30. SUE 32. 33. 34. 33. quinquella (Bedell) algeriensis sp. n. gilvipennella (Klimesch) comb. n. leucothorax sp. n. haraldi (Soffner) ilicis (Mendes) comb. n. heringella (Mariani) comb. n. alnifoliae sp. n. nigrosparsella (Klimesch) albifasciella complex (29—32) albifasciella (Heinemann) cerris (Zimmermann) montissancti (Skala) syn. n. pubescivora (Weber) comb. n. contorta sp. n. subbimaculella complex (33—36) subbimaculella (Haworth) nigrociliella (Stephens) syn. n. heringi (Toll) quercifoliae (Toll) sativella (Klimesch) syn. n. zimmermanni (Hering) syn. n. liechtensteini (Zimmermann) 36. phyllotomella (Klimesch) comb. n. 37. spec. (specimen 1375) terebinthivora group 38. terebinthivora (Klimesch) comb. n. “angulifasciella group” 39. erythrogenella (Joannis) 40. spiraeae Gregor & Povolny 41. agrimoniae (Frey) 42. hexapetalae (Szöcs) comb. n. angulıfasciella complex (43—46) 43. angulifasciella (Stainton) schleichiella (Frey) syn. n. utensis (Weber) syn. n. minorella (Zimmermann) syn. n. ? brunniella (Sauber) 44. atricollis (Stainton) aterrima (Wocke) staphyleae (Zimmermann) syn. n. 45. arcuatella (Herrich-Schaffer) 46. rubivora (Wocke) 47. spinosella (Joannis) 48. mahalebella (Klimesch) occultella group 49. occultella (Linnaeus) strigilella (Thunberg) ? mucidella (Hübner) mediofasciella (Haworth) syn. n. argentipedella (Zeller) 50. minimella (Zetterstedt) comb. n. mediofasciella auct. nec Haworth woolhopiella (Stainton) syn. n. viridicola (Weber) syn. n. Keys TO THE WESTERN PALAEARCTIC SPECIES OF ECTOEDEMIA SUBGENERA ZIMMERMANNIA AND ECTOEDEMIA S.STR. Based mainly on external characters!) 1. Forewings without distinct colour-pattern, irrorate or unicolorous, with at most incon- spicuous group of white scales at tornus.. 2 — Forewings with distinct white spot(s) or fascia): Hob ARR MONON ro Pate ODI, 12 . Frontal tuft dark fuscous brown to black 3 — Frontal tuft yellowish or orange, sometimes mixediw.ichituscousui m IR. 8 3. Thorax dorsally white with darker tips on N 1) Two species mentioned in the text, but still un- described have been excluded. 10 10. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 mesoscutum and tegulae. é with white hair-pencil 1. atrifrontella Thorax dorsally brown with at most white tips on mesoscutum and tegulae......... 4 . 6 hindwing without hair-pencil or costal emargination. ® with large patch of long tactile hairs on T7, extending almost to an- terior margin. Occurring in Afghanıstan ... N Gens ean oe ten tee 7. nuristanica dg hindwing with hair-pencil and usually costal emargination. If 2 with patch of long hairs, then only in posterior half of T7. Spe- cies occur in Europe or Anatolia . Hair-pencil in à short, about 1/4 hindwing length; without distinct costal emargination in hindwing. 2 unknown 4. hispanica Hair-pencil longer, at least 1/3 of hindwing length. Costal emargination conspicuous (isst ANT Re. 6 . Forewing with small tornal and costal white spots beyond middle, less conspicuous ın 3. Hair-pencil in d white. Forewing scales almost uniformly dark .... 2. hebwerdella Forewing with at most a tornal spot beyond middle. & hair-pencil fuscous or yellowish browne Sealestdarkenauepser ne 7 . 6 hair-pencil surrounded by brown scales. 2 with large patch of approximately 100 very long tactile hairs on T7 and 8 (visible without dissection). Only known from Greece and Anatolia 5. monemvasiae & hair-pencil surrounded by white scales. 2 with group of 20—30 long hairs, much shorter than in monemvasiae. Throughout Europe and Anatolia...... 3. longicaudella . Large species, forewing length 3.0—4.5 mm. Cilia-line indistinct. Aedeagus with 2 or 3 pairs of carinae. @ genitalia without vaginal sclerite Smaller species, forewing length 1.9—2.9 mm (rarely 3.0 mm). Cilia-line distinct. Ae- deagus with one pair of carinae only. 2 genitalia with vaginal sclerite .......... 10 . Ground colour dark brown, irrorate with white. d antennae with 36—41, ® with 36—37 segments. d hindwing with hair- pencil and costal emargination .... 6. amanı Ground colour lighter, more yellowish brown, irrorate with white. d antennae with 43—48, ® with 39—44 segments.d hindwing without hair-pencil or costal emargination 8. liguricella Ground colour white, with scattered brown Il, 12: 16. 17. 18. NEL . Scape white, with brown scales Ground colour brown to yellowish brown, mixed with yellowish white scales. & with- out hair-pencil, but with costal bristles (d of alnifoliae unknown) Scape white, without brown scales. Fore- wing with many yellow scales between brown ones. ® ductus spermathecae with 13—14 convolutions. Larva feeds on decid- uous Quercus in Europe . 28. nigrosparsella Scape white with some brown scales. Fore- wing mainly brown with few white scales. ® ductus spermathecae with 3 convolu- tions. Larva on Quercus alnifolia in Cyprus 27. alnifoliae (& unknown) Forewing with dorsal (tornal) spot only, but occasionally a few white scales along COSTA LE ent EE 13 FASCIATA it SAR 17 . Dorsal spot postmedial in position ..... 14 ° Dorsal spot medial in position ......... 15 Sc ae UE E. (Fomoria) or Trifurcula spp. Scape unicolorous white see 2 . Scales of forewing not significantly lighter at bases. Flagellum yellowish orange, simi- lar to scape and pedicel. & with hair-pencil, 2 with pointed ovipositor .... 9. intimella Scales of forewing distinctly lighter at base. Flagellum darker than scape and pedicel. d with costal bristles, 2 with blunt ovipositor ema dn ie on REE 16 Forewing with dorsal spot only. d hind- wing or forewing without androconial scales 25. ilicis Forewing usually with some white scales along costa, opposite dorsal spot, but not forming distinct spot. d hindwing upper- side and forewing underside with elongate patch of brown androconial scales......... 26. heringella Forewing with dorsal and costal distinctly postmediallin' position re ROL Aaa see 6, 2. liebwerdella Forewing with costal and dorsal spot medi- al or more basal, or fascia present ...... 18 Moth almost completely jet-black or grey- ish black, including cilia; cilia-line absent. Medial fascia present. Larva feeds on Betu- lacca Deere eee 19 Moth not completely black, usually with a fuscous or brownish tinge, cilia silvery white beyond distinct cilia-line. Larva feeds Onlotheritoodplantsirmi ee 20 3 underside of forewing with small patch 20. 22. 25} 24. 25} VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 11 of narrow white scales (difficult to see, fig. 86). Hair-pencil white. © frontal tuft yel- low or yellowish orange. Aedeagus without - long cornuti (fig. 405) 49. occultella d underside of forewing without small patch of narrow white scales, hair-pencil grey. ® frontal tuft black, occasionally mixed with some fuscous and/or yellow scales. Aedeagus with group of about 20 long cornuti (fig. 406) 50. minimella Forewing with white spots in addition to costal and dorsal spots or fascia ........ 21 Forewing with either fascia or costal and dorsal spot only 27 . Forewing with discal spot beyond middle. Brontaltuitiuscous black as daa 22.2. 22 Forewing without discal spot in second half. Frontal tuft usually with at least some yellow scales, but occasionally dark .... 23 Thorax completely white. Forewing with basal spot (sometimes small). 2 with group of long hairs on tergites 7 and 8. Feeds on evergreen Quercus......... 21. algeriensis Thorax fuscous black, with at most a white distal half. Forewing without basal spot, but sometimes with some white scales. ? with only few long hairs on T7 and 8. Feeds on deciduous Quercus...... 20. quinquella Forewings with many white scales scattered in basal half, sometimes becoming a discal spot or even confluent with other spots . 24 Forewings with only a basal spot or basal- dorsal streak, scattered white scales absent Ggpractically iso? St MORE RENT. 25 Frontal tuft yellowish orange to light ferru- ginous, never with fuscous scales. © with blunt ovipositor. & genitalia: valva without pointed tip (fig. 241), aedeagus fig. 361 N Ie) e Dt 10. bannoverella Frontal tuft light yellowish or yellowish fuscous to dark fuscous, never orange (light-headed 3 can not always be identi- fied with certainty on externals). © with pointed ovipositor (visible without dissec- tion). d genitalia: valva with pointed tip (fig. 242), aedeagus fig. 362.. 11. turbidella Costal spot distinctly more proximal than dorsal spot, not forming a fascia. Basal spot clearly separate from dorsal. d with costal bristles. without patch of long tactile hairs on T7 and 8 33—36. subbimacullela-complex Costal spot opposite dorsal, usually form- ing a fascia. Basal spot extending along dor- sal margin, often confluent with fascia. d 26. Li 28. 29. 30. 31. with hair-pencil or costal bristles. 2 with patch of many long tactile hairs on T7 and 8 Ee PCAN DETTE CALNE FR BARNA EA 26 Thorax white. Frontal tuft intensively orange. d with costal bristles. 2 terminalia complex, with thickened anterior apo- physes (fig. 444) .......... 23. leucothorax Thorax fuscous black. Frontal tuft yellow- ish, or mixed with fuscous. d with hair- pencil. Anterior apophyses not especially thickened (fig. 436) 15. caradjaı Fascıa or spots shining metallic silver (feed allkonvNosaceac I aa oa 38 Fascia or spots dull white or yellowish white (various foodplants, including Rosa- Cedo A Dati Sl Dem ects a! 28 Costal and tornal spot opposite, often forming fascia. d hindwing without costal bristles, in some species with hair-pencil 30 Dorsal spot distinctly beyond costal spot, usually not forming a fascia. d hindwing with costal bristles, hair-pencil absent .. 29 Thorax usually uniform dark. Forewing ground colour almost uniformly blackish ern scales only slightly lighter at bases. 3 aedeagus with two pairs of carinae, valva fig. 244. 2 bursa with pectinations. Larva on Ulmus 14. preisseckeri Thorax usually with white tips of mesoscu- tum and tegulae. Forewing ground colour fuscous blackish, slightly speckled because of lighter scale bases. d aedeagus with one pair of carinae, valva figs. 261—264. 2 bur- sa without pectinations. Larva on decidu- ous Quercus 29—32. albifasciella-complex Thorax with or without white tips. Fore- wing ground colour brown, more irrorate than preceding species, scales, especially at forewing tip only dark at their tips. d ae- deagus with one pair of carinae, valva fig. 255. 2 bursa without pectinations. Larva on evergreen Quercus 24. haraldi Basal half of forewing with scattered white Selen tr ne er see 24 Basal half of forewing never with white scaleswoursidemheispotste an ne 31 Large species, forewing length 2.6—3.2 mm. Antennae in d with 49—60 segments, in 2 with 34—39. & with hair-pencil, never with brown lamellar androconial scales . 32 Smaller species, forewing length 1.7—2.5 mm. Antennae in 6 with 30-40, in 2 with 23—35 segments. d with or without hair- pencil, with or without brown androconial SCHES veers ine 19, ARE anc elite 35 DDR 33. Dn 93. 36. DI 38. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Costal and dorsal spot forming distinct fas- cia. d with white or brown hair-pencil. © with broad oval signa of same length. Medi- - terranean species, feed on evergreen Quer- cus DI) Costal and dorsal spot clearly separate. ó with yellow hair-pencil. 2 with elongate signa of different length. European species, feed on Populus 34 3 hair-pencil white. Capsule length 260— 300 um. 2 with dense patch of very long tactile hairs on T7 and 8 (fig. 437) EDER O BERN 17. suberis dg hair-pencil ochreous-brown. Capsule length 220—260 um. 2 without long tactile hairs on T7 and 8 (fig. 438) .. 18. andalusiae 3, or with 34—38 antennal segments. On P. alba 12. klimeschi 2 only, parthenogenetic, with 26—32 an- tennal segments. On P. tremula........... 13. argyropeza d with patch of brown androconial scales on upperside of hindwing and underside of forewing. Forewings ochreous brown, or greyish brown with yellowish tinge, com- paratively light. Fascia ill-defined. Mediter- GANCANISPECIE EEE RR: 36 3 without brown androconial scales. Fore- wings definitely dark, fuscous black. Fascia distinct. Species from central and southeast- ern Europe 37 Frontal tuft yellow, orange or fuscous. 6 without hair-pencil. 2 genitalia with elon- gate signa (fig. 202) 38. terebinthivora Frontal tuft yellowish white. d with yel- lowish white hair-pencil. 2 genitalia with small oval signa (fig. 176) . 19. aegilopidella Very small species, forewing length 1.7— 2.1 mm. d without hair-pencil. Aedeagus with spinose dorsal process and ventral ca- rinae. 2 genitalia with vaginal sclerite. Feeds on Filipendula vulgaris............. 42. hexapetalae Larger, forewing length 2.2—2.5 mm. & with yellowish white hair-pencil. Aedeagus without carinate processes. ® genitalia without vaginal sclerite. Feeds on Spiraea media 40. spiraeae Frontal tuft very dark, blackish fuscous. 3 hindwing with white hair-pencil 46. rubivora Frontal tuft varying from yellowish orange or pale ochreous to fuscous, but never black. d hindwing with or without hair- PEIN Sy payors ay) eal a ee 59 39! 40. All. 42. A9? 44. A5). 46. a Forewing with costal and dorsal spot usual- ly separate, dorsal spot distinctly beyond costal spot. Frontal tuft ferruginous, with sometimes fuscous scales on crown; collar yellowish white. 6 hindwing without hair- pencil Meine ae ee 39. erythrogenella Forewing with costal and dorsal spot often united to form constricted fascia. Dorsal part of fascia not distinctly beyond costal part. d hair-pencil present or absent.... 40 dui na ehe 41 SM; 45 Hindwing with fuscous hair-pencil, sur- rounded by patch of brown scales. Small species, forewing length 1.4—2.1mm...... 47. spinosella Hindwing with white hair-pencil 42 Hindwing without hair-pencil......... Collar yellowish orange to ferruginous, ap- proximately same colour as frontal tuft. Valvae with inner margin distinctly sinuate dele Ge ele ere pe 43. angulifasciella Collar brown to black, darker than frontal tuft. Valvae with inner margin approxi- mately straight maa 00 NC SRI Frontal tuft ferruginous yellow, often mixed with fuscous. Smaller species, fore- wing length 1.8—2.3 mm. Feeds on Fraga- ria and Potentilla 46. arcuatella Frontal tuft orange to ferruginous. Slightly larger, forewing length 2.2—2.7 mm. Feeds on Rosaceous trees and Staphylea ot. A 44. atricollis Frontal tuft yellowish to ferruginous, or even fuscous. Collar greyish brown. Scape often with brown scales. Forewing length 2.3—3.0 mm. Tegumen pointed. Feeds on ARRONE 41. agrimoniae Frontal tuft and collar yellowish orange to ferruginous. Scape uniform white. Fore- wing length 1.9—2.4 mm. Tegumen round- ed. Feeds on Prunus spp... 48. mahalebella Collar and frontal tuft concolorous, yel- lowish orange to ferruginous 46 Collar distinctly different in colour from frontal tuft: greyish brown to fuscous black. Frontal tuft yellowish orange to fus- cous Larger species, forewing length 2.0—2.9 mm. Signa elongate (figs. 211, 212) VIGO ALA 43. angulifasciella Smaller species, forewing length 1.9—2.4 aon, SMe Ovvell Gul, BN QMO) os so caacc vet bethere ns Sede 48. mahalebella Scape usually with some brown scales, es- 48. 49. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 13 pecially along distal margin. Collar greyish brown. Ovipositor slightly pointed........ Has a ae ur 41. agrimoniae Scape uniform white. Collar fuscous to blacks Ovipositor blunt. aa... nino. 48 Small or medium sized species, forewing length 1.4—2.4 mm. Signa length 200—370 ED 46 à Se RER ER ee EE 49 Larger species, forewing length 2.3—2.8 mm. Signa distinctly longer, 380—490 um. Feeds on Rosaceous trees and Staphylea.... BPE HRA se send: HEURE 44. atricollis Small species, forewing length 1.4—2.1 mm. Signa with smooth, uniformly curved outline, longest 250—370 um, shortest 230—330 um, 2.4—3.5 X as long as wide. Feeds on Prunus spp......... 47. spinosella Medium sized species, forewing length 1.7—2.4 mm. Signa with irregular outline, longest 230—310 um, shortest 205—280 um, 3.1—4.1 X as long as wide. Feeds on Fragaria or Potentilla 45. arcuatella Based mainly on male genitalia!) . Aedeagus with three pairs (two in liguricel- la) of carinae, with ventral pair usually very prominent and longer than other carinae; dorsal carinae sometimes composed of sev- eral spines (palmate). Valva: tip straight or only very slightly curved inwards; often an inner (mesal) lobe present; large genitalia, capsule 320—430 um, aedeagus 370—500 um. (350 in nuristanica); valva longer than 270 um. Subgenus Zimmermannia ...... 2 Aedeagus with one or two pairs of carinae, or carinae absent, ventral pair not more pronounced and larger than dorsolateral pair. Valva tip usually curved inwards, in- ner lobe absent. Genitalia usually smaller, capsule 150—320 um (—390 in occultella); aedeagus 205—410 um; valva shorter than 270 um (except leucothorax and occultella, to 320 um). Subgenus Ectoedemia ....... 9 . Aedeagus with two pairs of carinae, the ventral pointed and widely separate. Aedea- gus with two distal spinose lobes (figs. 356—358). Valva (fig. 238) with inner lobe. Tegumen with tongue shaped process (fig. 336) 8. liguricella Aedeagus with three pairs of carinae, the ventral not widely separate, aedeagus with- 1) Males of E. alnifoliae are unknown. 10. out spinose lobes. Tegumen without tong- Wea ped | PROCESS an ne CURE. 3 . Capsule very wide (+ 370 um), almost as wide as long. Valva broad (fig. 236). Aedea- gus almost half as wide as long, dorsal cari- nae comprising row of 4—5 teeth 6. amani Capsule narrower, less than 360 um wide. Valva narrower, aedeagus less than half as wide as long. Dorsal carinae simple, bifid or Palmare mente hs vee ace sale cn 4 . Ventral carinae prominent, long and point- ed, larger than dorsal and lateral pairs. Hindwing with hair-pencil............. 5 Ventral carinae about same size as dorsal and lateral, with bifurcate tip. Valva nar- rowed before tip (fig. 237). Hindwing with- out hair-pencil 7. nuristanica . Valva with prominent inner lobe, approxi- mately, amma logan eras rere nee tee 6 Valva without inner lobe, or with very slight lobe, not projecting beyond inner Maine, Bean nen Med. 7 . Gnathos with narrow pointed central el- ement. Aedeagus with palmate dorsal cari- nae and stout triangular cornutus SE hae ttes Set o] 5. monemvasiae Gnathos with broad triangular central el- ement. Aedeagus with single or bifurcate dorsal carinae, without stout cornutus ..... 4. hispanica . Aedeagus clearly constricted in middle. Dorsal and lateral carinae connected by dis- tiNnctiti eens ira 8 Aedeagus not constricted in middle. Dorsal and lateral carinae not connected by rim.... 3. longicaudella . Outer margin of ventral carinae distinctly serrate. Tip of valva rounded. Ventral arm of transtillae very short .… 1. atrifrontella Outer margin of ventral carinae smooth or with a few spines. Tip of valva always slightly hooked. Ventral arm of transtilla usuallylongern. ran. 2. hebwerdella . Aedeagus with ventral and dorso-lateral ca- OON a Blo Re chon Des Van tee eee LE Lio eae re 10 Aedeagus with ventral carinae only, sometimes divided, or noneatall....... 14 Dorso-lateral carinae stout, curved in lateral view, larger than ventral pair, often bifurcate. Ventral carinae connected by bas- allplareh Sp ire 11 Dorso-lateral carınae same size as ventral carinae or smaller, not particularly stout. Ventral carinae not similarly connected . 12 . Valva ending in abruptly narrowed tip. Ae- 14 12, 5). 16. 17% 18. 19. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 deagus not markedly asymmetrical (fig. 362) 11. turbidella Valva gradually narrowing towards up. Ae- deagus markedly asymmetrical (figs. 363, 400, 401) 12. klimeschi Ventral carinae curved, often overlapping (fig. 364). Gnathos triangular, pointed ERNEUTE EM 14. preisseckeri Ventral carinae straight, distinctly separate. Gnathos with rounded central element. . 13 . Valva with broad blunt tip, widest beyond middle. Gnathos with spines on central el- MICA I I Se 10. hannoverella Valva with pointed tip, widest at basis. Gnathos broad, without spines 9. intimella . Tegumen cuspidate, long pointed. Carinae divided each in at least 4 similar spines (figs. BOARDS) ET END MALE PRE 15 Tegumen triangular, rounded or blunt. Ca- rinae single, or with some additional, usual- ly smaller, spines Small species, aedeagus 180—230 um. Cari- nae distinctly below tip of aedeagus, with pointed tips. Ventral process with some spines. Gnathos triangular . 41. agrimoniae Large species, aedeagus at least 280 um long. Carinae with blunt tips reaching tip of aedeagus. Ventral process smooth. Gnathos blunt ge wa bl 16 Aedeagus with small triangular cornuti only (fig. 405). Gnathos with very wide, blunt, central element (fig. 327). Comparatively large, aedeagus 305— 350 um . 49. occultella Aedeagus with a row of about 20 long elon- gate cornuti at right side (fig. 406). Gnathos with narrow, truncate, central element (fig. 328). Smaller species, aedeagus 280—310 um 50. minimella Aedeagus without any carinae. Tegumen protruding, triangular (fig. 127). Gnathos with central element reduced (fig. 318). Valva fig. 271 40. spiraeae Aedeagus with ventral carinae. If tegumen protruding, than blunt or rounded. Gna- thos usually with distinct central element ERE U GEO ARE RED ame 18 Aedeagus dorsally with medial spinose pro- cess (fig. 403, 404). Small species, capsule 195—240 um, as wide as long, and aedeagus relatively long, 260—290 um 42. hexapetalae Aedeagus without dorsal spinose process 19 Capsule 150—170 um long, wider than long. Tegumen bulbous (fig. 410). Gnathos 20. ZA DDR 257 DAS 258 with central element in form of transverse bar (fig. 298) 19. aegilopidella Capsule longer than 190 um. Tegumen not bulbous. Gnathos with central element not In LOSMVOL trans Versel ate see Carinae with many small additional spines at base (figs. 28, 390—393). Tegumen prominent, longer than wide, cut off (fig. 413). Valva with inner margin straight or slightly sinuous Carinae without or with few additional spines. Tegumen triangular, rounded or wider thanilongan urn... en Valva with inner margin sinuous, forming a slight bulge beyond middle (fig. 273). Cap- sule length 210—260 um, aedeagus 215— 275 um 43. angulifasciella Valva with inner margin straight below apex (figs. 274276) 44. atricollis (capsule 270—290 um, ae- gedagus 260—290 um, head orange) 46. rubivora (capsule 255—285 um, aedea- gus 235—265 um, head black) 45. arcuatella (capsule 250—255 um, ae- deagus 230—245 um, head orange) Valva with many setae on dorsal and inner surface, the prominent sockets result in a distinctly serrate inner margin (figs. 260— 266). Gnathos undivided, without serrate margins. (Hindwing with costal bristles) 23 Valva with comparatively few setae on in- ner and dorsal surface, usually restricted to posterior half; rarely a few prominent sock- ets along inner margin, never distinctly ser- rate. Gnathos divided or undivided, with or without serrate margins. (Hindwing with or without hair-pencil or costal bristles) 24 Gnathos with central element truncate (figs. 307—312). Aedeagus with carinae simple... 29—32. albifasciella-complex (forewing with white spots) 28. nigrosparsella (forewing irrorate) Gnathos with central element rounded (figs. 313—315). Aedeagus with carinae usually with a few additional spines ....... 33-36. subbimaculella-complex Tegumen wider than long, truncate (fig. 412). Gnathos with very short central el- ement. Valva fig. 268 .... 38. terebinthivora Tegumen longer than wide, or not protrud- ing at all. Gnathos with conspicuous central clement NR 25 Valva with inner margin almost straight up to the distinctly separate apex. Small spe- cies, capsule length 190—225 um... 26 26. 27. 28. 29. 30. Van NIEUKERKEN: Western Palaearctic Zimmermannıa and Ectoedemia 15 Valva with inner margin markedly concave, especially in distal half, with gradual tran- sition into apex. Small or large species .. 27 Gnathos with smooth triangular central el- ement. Valva with apex almost posteriorly pointing (fig. 269) 39. erythrogenella Gnathos divided, basal part with more or less serrate margin. Valva with inwards GUvediapex (fest 277,278). man ala... BAU. ei; 47. spinosella (with hair-pencil) 48. mahalebella (without hair-pencil) Gnathos with smooth, undivided triangular or tongue-shaped central element. Aedea- gus distinctly longer than capsule, carinae simple 28 Gnathos with divided central element, basal part with serrate margins, distal part spatu- late. Aedeagus about as long as capsule, shorter or slightly longer. Carinae usually withladchtionalspineser nnn el 31 Valva with inner margin basally convex, apically concave, with sharp delimitation. Aedeagus very long, 310—395 um... 29 Valva with inner margin basally hardly con- vex, without sharp delimitation between basal part and concave distal part. Aedeagus shorter, 275—290 um (305 in specimen 1843) 30 Tegumen produced in broadly triangular pseuduncus with rounded tip. Capsule lon- ger than 260 um 17. suberis Tegumen broad, truncate, not produced in- to pseuduncus. Capsule 225—235 um... 18. andalusiae Tegumen produced into distinct rounded pseuduncus. Aedeagus 275—290 um....... 15. caradjaı Tegumen truncate, not produced into pseu- duneus aedeious 305 um ne see ee le 16. specimen 1843 . Valva with very prominent bulging outer margin (122255) Ae ee an. 24. haraldı Valva with outer margin uniformly convex 32 . Valva dorsal surface with back-folded lobe (fig. 253). Hindwing with prominent black hair-pencil 22. gilvipennella Valva without dorsal lobe. Hindwing with lighter hair-pencil or without.......... 55 . Valva extremely long and narrow (fig. 254), longer than 270 um. Aedeagus distinctly shorter than capsule... 23. leucothorax Valva not extremely long and narrow, shorter than 260, usually shorter than 220 34. um. Aedeagus as long as capsule or longer DE SEDIA ASIA EERIE). SRE AE Le. 34 Hindwing with costal bristles. Forewing withädorsalispotonly. 22.2.2. see 35 Hindwing with hair-pencil. Forewing with at least three spots 36 . Hindwing upperside and forewing under- side with brown androconial scales 26. heringella 25. ilicis . Tip of valva pointed. Hair-pencil yellowish 20. quinquella Tip of valva truncate. Hair-pencil white .... 21. cf algeriensis Based mainly on female genitalia!) . Corpus bursae longer than 880 um, usually longer than 1000 um. Vestibulum with in- conspicuous sclerotisations or spines, with- out vaginal sclerite. Margin of signa wider than individual cells. Subgenus Zimmer- NE NS oes Be re 2 Corpus bursae usually shorter than 880 um, but occasionally up to 935 um, and then al- ways with vaginal sclerite. Margin of signa narrower than individual cells. Subgenus Betoedenna ash. een sie 6 . Ductus spermathecae with 12/—13 con- volutions.Vestibulum with two groups of SPIIEST Rn Ie FR 6. amanti Ductus spermathecae with 412—5V4 con- vo lutions de eeen ee 3 Ductus spermathecae with 2/—3% con- volutionsai Aat gan ee 5 . T7 with large patch of long tactile hairs, ex- tending almost to anterior margin (fig. 426) ef TREND IE ne ef 7. nuristanica T7 with long hairs only at posterior margin . T7 and 8 with dense bunch of many long hairs (fig 424). Longest signum longer than 500 um. Eastern mediterranean species..... at era Me 5. monemvasıae T7 and 8 with some long setae in a row, not forming a dense bunch (figs. 427, 428). Longest sıgnum shorter than 500 um. West- ern mediterranean species..... 8. liguricella . Ductus spermathecae with 3/—3% con- VolUtionsé bikie? mnd 3. longicaudella Ductus spermathecae with 2'2—3 convolu- 1. atrifrontella or 2. hebwerdella ') Females of E. hispanica are unknown. 16 il. 10% 157 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 . Corpus bursae without pectinations, com- pletely smooth outside signa. A group of densely packed pectinations present in ves- (LEA UE PR a Cee RON NE 7 Corpus bursae mostly covered with small pectinations or spines. Densely packed pec- tinations in vestibulum may be either pre- sent or absent 14 . Signa of about same length, approximately 2.6—3.4 X as long as wide. Anterior apo- physes much widened (fig. 444). Large group of many long hairs along posterior MATOS ae 23. leucothorax Signa distinctly dissimilar, more than 3.5 x as long as wide. Anterior apophyses not markediyawidenedwr ee 8 . Ductus spermathecae with more than 31% Convolutionsmas m ae m. 9 Ductus spermathecae with 2—3V4 convolu- ONSEN RUES i OR ten coil: 10 3/2—4 convolutions ........... 30. cerris M eee 31. pubescivora 10Y%2—12 (rarely 1312) convolutions VEN EEE IE 32. contorta 1312—14 convolutions .. 28. nigrosparsella . T7 and 8 with in total more than 70 setae, including some very long (fig. 441). Anal papillae each with more than 24 setae lara, Beer zahl 21. algeriensis T7 and 8 with much fewer setae, usually not exceeding 25. Anal papillae with less than ZA SERA CAMPI WIRE ARCS. bo REN 11 Abdominal tip narrow, T8 and 9 with dis- tinctly converging margins (fig. 429). Spicu- late pouch with small, single denticles, ap- proximately equally spaced ... 9. intimella Abdominal tip wider, T8 and 9 not so dis- tinctly converging. Spiculate pouch with small denticles, often in small groups, not equallysspacedae nen ge RR 12 T7 with distinct row of 6—14 setae along Posten opm an 13 T7 without such a row, at most few scat- teredishoraserae sro ra ate 24. haraldi, 20. quinquella or 27. alnıfoliae, compare externals and figures of female ter- minalia (figs. 445, 440, 449). Sides of S8 almost parallel. Convolutions of ductus spermathecae very wide (fig. 416) ... 29. albifasciella Sides of S8 diverging anteriorly. Convolu- tions of ductus spermathecae narrow (figs. AIS) ane NN TES al 22. gilvipennella, 25. ilicis, 26. heringella or ry 14. Sy. IO, 117 18. 19. 20. De 33—36. subbimaculella-complex, compare externals and figures of female terminalia. T7 and 8 covered with many (more than 100) very long hairs, reaching abdominal tip. Ductus spermathecae with 3/— 4/5 convolutions. ERRORE 15 T7 and 8 covered with few short setae only, at most 20. Ductus spermathecae either with 5 or less than 31/2 convolutions... 16 Bursa almost globular. Long setae smooth. Ductus spermathecae with 4—4 distinct convolutions. Abdominal tip fig. 437 17. suberis Bursa elongate. Long setae pectinate. Duc- tus spermathecae with 3!2—4 less distinct convolutions. Abdominal tip fig. 436 15. caradjai Ductus spermathecae with 5/2 convolu- 18. andalusiae Ductus spermathecae with 2—3'/ convolu- | 17 Vestibulum with patch of densely packed pectinations near entrance to ductus sper- mathecae. Vaginal sclerite present. Spiculate pouch conspicuous, usually with many dis- tinct spines (sun. AIO ee eee 18 Vestibulum without patch of densely pack- ed pectinations. Vaginal sclerite present or absent. Spiculate pouch inconspicuous or absent 23 Signa completely different in form and length, longest reaching into vestibulum, shortest 4.2—5.0 X as long as wide. Termi- nalia fig. 459 38. terebinthivora Signa similar in form, sometimes slightly different in length, not reaching into vesti- bulum 19 T7 with distinct row of 4—12 setae along posterior margin. Spines of spiculate pouch: not all equally spaced, or very few only . 20 T7 without distinct row of setae along pos- terior margin. Spines of spiculate pouch dis- tinct, all equally spaced, not grouped ... 21 Signa 2.4—3.0 X as long as wide. Terminal segments narrow, fig. 435 .. 14. preisseckeri Signa 3.0—5.6 X as long as wide. Terminal segments wider, fig. 460 39. erythrogenella T8 and 9 posteriorly narrowed, forming pointed ovipositor. Posterior apophyses widened at anterior tips (figs. 431, 432). .... else mp e 11. turbidella T8 and 9 not so much narrowed, ovipositor blunt. Posterior apophyses not distinctly widened 22 VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 17 22. T8 about 2 X as wide as long (fig. 430). Anal papillae with 6—11 setae. Signa 390— 480 um. S8 without pronounced corners ... EN EN 10. hannoverella — TS more than 2 X as wide as long (fig. 433). Anal papillae with 9—11 setae. Signa 373— 416 um. S8 with pronounced corners ...... LARMES 12. klimeschi — T8 more than 2 x as wide as long (fig. 434). Anal papillae with 5—9 setae. Signa 270— 394 um long. S8 without pronounced cor- MERS pode SA ae 13. argyropeza 23. Anal papillae with 18—40 setae. Terminal segments wide (figs. 470, 471). Pectinations in bursa in two longitudinal bands, running halfway between the signa 49. occultella or NN EMF REC 50. minimella — Anal papillae with 4—16 setae. Terminal segments not so wide. Pectinations in bursa more regularly distributed ............ 24 24. Vestibulum completely smooth, without vaginal sclerite or spiculate pouch. T8 di- idedinimiddle. men pesa ann ae. 25 — Vestibulum with vaginal sclerite, although sometimes indistinct, and sometimes with inconspicuous spiculate pouch. T8 undi- leder ingenieur dida 26 25. Signa dissimilar, 320—440 um long. Anal papillae with 13—16 setae. Terminal seg- MANS REL NEON. ent 40. spiraeae — Signa similar, 180—300 um long. Anal pa- pillae with 7—12 setae. Terminal segments fig. 462 41. agrimoniae 26. Bursa very small, 310—350 um. Signa short, 189—223 um, oval, occupying large EME Or DUR EP RE 19. aegilopidella — Bursa larger, 570—715 um. Signa short, 200—300 um, oval, confined to posterior [afosa nn). 48. mahalebella — Bursa intermediate, 400—660 um. Signa variable in length, elongate, throughout Dui SB REED. Ra N RE ce sit 42. hexapetalae, 47. spinosella and 43—46. angulıfasciella-complex, compare externals, diagnoses, or genitalia figures. Subgenus Zimmermannia Hering Zimmermannia Hering, 1940: 266. Type-species: Ectoedemia liebwerdella Zimmermann, 1940, by original designation and monotypy. Ectoedemia sensu Klimesch, 1953: 163 [European species]. Ectoedemia (Zimmermannia); Schönherr, 1958: 6: Borkowski, 1972: 699; Emmet, 1976: 188, 203. Ectoedemia castaneae group sensu Wilkinson & Newton, 1981: 72. Description. Adult. Relatively large nepticulid moths, forewing length 2.8—4.5 mm, wingspan 6.4— 9.8 mm (in Palaearctic species). Head. Antennae long, more than half length of forewing, in male with 36—58 segments, in female with 36—49 segments. Scape and pedicel white, flagellum darker. Wings. Uniform irrorate ochreous or yellow- ish-white, with darker scaling, often predomi- nantly brown, without fascia, sometimes a small dorsal (tornal) and/or costal spot present. Cilia- line not distinct. Hindwing in male without cos- tal bristles, hair-pencil present in most species, surrounded by special scales. Humeral lobe of- ten prominent, beyond which hindwing is sud- denly emarginated (figs. 10—14). Forewing venation (fig. 8). R and M + Cu forming closed cell, branches R,, R,,;, Ry, Rs, M and Cu present. A thickened, without anal loop. Cu and A often very long, seeming fused at tips. Male genitalia. Vinculum ring-shaped, ante- rior extension not long, anteriorly convex. Te- gumen slightly produced into a triangular or blunt pseuduncus. Uncus absent. Gnathos with prominent spatulate or triangular central el- ement, margins smooth. Valva approximately triangular, tip not separate, usually not curved inwards; often with a mesal (inner) lobe. Aedea- gus stout, with large ventral carinae, smaller dorso-lateral carinae and usually dorsal carinae. Ventral carinae fitting by membranes to fold in dorsal surface of valvae. Dorsal carinae palmate in some species. Vesica with numerous denticu- late cornuti and usually one large cornutus or sclerotised plate posteriorly. Female genitalia. On tergites 7 and 8, near an- terior margin of T8 usually a group or row of very long setae, T8 with many shorter setae, without scales. Anal papillae with setae. Poste- rior apophyses often reaching beyond anterior apophyses. Vestibulum with indistinct paired sclerotisation, or with groups of spines, vaginal sclerite or spiculate pouch absent. Corpus bur- sae long, elongate, covered with pectinations, except in anterior part, arranged in concentric bands around long signa. Margin of signa wider than individual cells. Larva. Long yellow larvae with strongly scle- rotised head-capsule, feeding venter upwards. 18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 See Schönherr (1958) and van Nieukerken & Jansen (in preparation). E. lebwerdella has six to eight larval instars (Schönherr, 1958). Biology. The larvae of the species where the life histo- ry is known, are bark-miners (or gall-makers in bark: E. castaneae) in Fagaceae (Fagus, Quer- cus, Castanea), and Ulmaceae (E. amani only). The mines are galleries. The larvae feed for one or two years and leave the mine in spring to pu- pate in the soil. Adults fly throughout the sum- mer. The life-history became particularly well- known through the excellent work of Schönherr (1958) on E. hebwerdella, and the studies of Busck (1913, 1914a,b) on American species. Distribution and composition. Eight species are described here from the Western Palaearctic region as far east as Af- ghanistan, two species (E. admiranda and E. sı- vickisi) were described by Puplesis (1984b) from the Eastern Palaearctic region, and twelve spe- cies were recorded from North America by Wilkinson & Newton (1981) and Wilkinson (1981), and more unnamed Nearctic species are present in collections. Remarks. The species of this subgenus are remarkably uniform both in external features and genitalia, also when the Nearctic species are taken into consideration. Although the life history of only 3 Nearctic and 4 Palaearctic species is (partly) known, it seems very probable that all species are bark-miners, and the majority feeds on Fa- gaceae. Several of the species of which the life- history is unknown have also been collected in vegetation containing many Fagaceae (Quer- cus). There are mines also known which have not yet been associated with existing species. Schonherr (1958) for instance reported mines on Carpinus, and he and Klimesch (1953) on Cas- tanea. I also found mines on Castanea and Quercus ilex in the south of France, and on Q. coccifera in Spain. Unfortunately rearing of Zimmermannia larvae proved very difficult, so it will probably be a long time before the life- histories of all species have been worked out. 1. Ectoedemia (Zimmermannia) atrifrontella (Stainton, 1851) (figs. 8, 10, 33, 34, 35, 89, 143, 144, 231, 281, 3295 337) 93989 246 1215072514) Trifurcula atrifrontella Stainton, 1851: 11. 2 Syntypes, England, G. Bedell (depository unknown) [not examined]. Zimmermannia heringiella Doets, 1947: 504—506, 5 figs. Lectotype d (here designated), Netherlands: Hollandse Rading, 15.v11.1946, el. Quercus, J. Doets, Genitalia slide V. 679 on pin (RMNH) [ex- amined, genitalia figured by Doets]. [Synony- mised by Klimesch, 1953]. Trifurcula atrifrontella; Stainton, 1854: 306; Herrich- Schaffer, 1855: 360; Stainton, 1859: 438; Wocke, 1871: 335; 1874: 97; Heinemann & Wocke, 1877: 726; Meyrick, 1895: 727; Tutt, 1899: 358; Rebel, 1901: 221; Meess, 1910: 482; Meyrick, 1928: 864; Beirne, 1945: 207, 208; Gerasimov, 1952: 202; Karsholt & Nielsen, 1978: 3, 4, figs. 7, 8 (d geni- talia). Ectoedemia done: Klimesch, 1953: 191—193, fig. 18 (revision, 6 genitalia); 1961: 749; Lhomme, 1963: 1210; Szöcs, 1965: 49; Bradley et al., 1972: 3; Borkowski, 1975: 496; Emmet, 1976: 203 pl tiga, pl he Trifurcula (Ectoedemia) atrıfrontella; 1971: 245. Johansson, Diagnosis: the white thorax together with the black head separate atrıfrontella from other Zimmermannia species, the snow-white hair- pencil in the male is a good additional character separating it from longicaudella. The narrow capsule, constricted aedeagus, serrate carinae and short ventral arms of transtilla are diagnos- tic characters of the male genitalia. The female genitalia differ from longicaudella by shorter posterior apophyses and lower number of con- volutions in ductus spermathecae, but cannot be separated from liebwerdella. Description. Male (fig. 35). Forewing length 2.88—3.24 mm (3.05 + 0.13, 12), wingspan 6.5—7.4 mm. Head: frontal tuft and collar dark brown to black. Antennae long, with 45—53 segments (48.3 + 2.7, 7). Thorax yellowish white, except brown caudal tips of mesoscutum and tegulae. Forewings dark brown, irrorate with varying amount of white, tornal spot usually white; cilia silvery white beyond ill-defined cilia-line. Hindwing (fig. 10) with snow-white hair-pencil of approximately % hindwing length, sur- rounded by white lamellar scales; humeral lobe prominent, costal margin distinctly emarginated beyond hair-pencil. Female. Forewing length 3.2—3.84 mm (3.59 + 0.23, 9); wingspan 7.2—8.5 mm. Antennal segments 37/—49 (42.2 + 3.2, 10). Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 19 Male genitalia (figs. 89, 329). Capsule length 390—411 um (404.2 + 8.7, 7), slender, width 274—304 um. Vinculum with posterior part of ventral plate less than one third of ventral plate. Tegumen slightly cuspidate. Gnathos (fig. 281) with central element long and narrow, parallel- sided. Valva (fig. 231) slender, length 287—321 um (297.6 + 12.6, 7), approximately triangular, without any lobe along inner margin, tip round- ed; transtilla with ventral arm extremely short. Aedeagus (figs. 337, 338, 346) 450—501 um (471.4 + 19.2, 7), constricted at level of opening for ductus ejaculatorius; ventral carinae long, approximately one third of total length aedea- gus, with distinct serrate outer margins; lateral and dorsal carinae connected by prominent rim, stout and pointed, the dorsal longer. Vesica with distally a sclerotised plate with indistinct folds or ridges in addition to small cornuti. Female genitalia (figs. 33, 34, 143, 144, 421). T8 with many long hairs, more than 50, a row of 10—20 thicker and very long setae along an- terior margin of T8, scales absent; anterior mar- gin of T8 slightly indented. Anal papillae with 8—13 setae. Posterior apophyses hardly reach- ing beyond anterior apophyses. Vestibulum with pair of indistinct sclerotisations. Corpus bursae 1080—1270 um, covered with pectina- tions, partly in concentric bands around signa; signa elongate, similar, length 473—572 um (510 + 41, 14), 4.4—6.7 X as long as wide. Ductus spermathecae with 2'2—3 convolu- tions, becoming wider distally. Larva. Yellow, very elongate. Head-capsule brown. Ventral plates absent. Biology. Host plants: Quercus robur L., Q. pubescens Willd. and probably other Quercus species. In Spain the species was collected in cork-oak woods with some Quercus faginea Lam., of which the latter is the most likely foodplant here. Mine (fig. 472). Contorted gallery in smooth bark of branches and thin trunks. The larva feeds mainly in the direction of the main axis. Life history. Incompletely known, larvae start feeding probably in summer and over- winter at least once, but analogous to liebwer- della and longicaudella it could have a two year cycle. Full grown larvae collected late May and June pupate soon and emerge within a few weeks. Adults are frequently caught at light from early July until the middle of September. Rearing is difficult, and actually very few speci- mens have been reared. Distribution (fig. 514). Widely distributed in Europe from southern Finland to Spain, but not recorded from eastern Europe, except Hungary, nor from Belgium (Janmoulle’s 1947 record actually refers to longicaudella), Ireland, Norway or Portugal. This is the only Zimmermannia species known from Great Britain. In central and southern Eu- rope this species is often less common than longicaudella. Occurrence in Anatolia (one un- certain female) has to be confirmed. Remarks. Stainton described this species from two specimens from Bedell’s collection. Unfortu- nately these specimens could not be found in BMNH, and the collection seems to have been dispersed after auctioning, so the types remain unknown. The identity of this species however seems to be beyond doubt, since there are two subsequent correctly identified specimens in Staintons collection, which represent this, the only British Zimmermannia species. Records prior to 1953, and also several more recent ones, cannot be relied on since they refer at least partly to E. longicaudella. The life-history of this species was discovered by Doets (1947), who at that time described it as the new species Zimmermannia heringiella. Previously E. atrifrontella was incorrectly be- lieved to mine bark of Sarothamnus. Material examined: 26 6, 18 2. — Austria: 5 6, Gumpoldskirchen, Glaslauterriegel, 10.vı1.1958, 10.v11.1981, 26.viu.1983, and 1.1x.1983, F. Kasy; 1 6, Hundsheimer Berg, Porta Hungarica (near Hain- burg), 2.vu.1977, F. Kasy (NMW). — France: 1 d, “Antarv”. (? near Digne), 13.vi11.1903, Chrétien; 1 d, Digne, vı1.1903, Chrétien (MNHN); 2 dg, Viens (Vaucluse) (near Apt), 6.vı1.1974, 1.1x.1975, Buvat (coll. Buvat). — Germany, West: 2 ©, Leine, Eime, 10.viii.1889, coll. J. Schlumberger. — Germany, East: 1 2, Altenburg, Krause (MNHN). — Great Britain: 1 2, Dartford Heath (Kent), 12.vin.1892, Tyerman; 1 3, Ham Street (Kent), 16.1x.1961, S. Wakeley (UMZC); 1 2, Lewisham (London), 13.vm.1851, beaten from oak, J. Stainton; 1 d, 1851, J. Grant, no further data (BMNH); 1 à, 1897, J. B. Hodgkinson, no further data; 2 d, no data, Whittle coll. (genitalia figured by Klimesch, 1953) (BMNH). — Hungary: 1 2, Nadap (near Velencei-t6), 6.1x.1951, Kovacs (TMAB). —Netherlands: 3 d, 4 ©, Hilversum, el. 10—17.vii.1948, el. 21.viii.1950, Quercus, Doets 20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 (RMNH, 1 MHUB); 1 6, 3 ® (lecto- and paralecto- types of heringiella Doets), Hollandse Rading, e.l. 10—15.v111.1946, Quercus, Doets (RMNH, ZMA); 1 d, Leuvenum, Ullerberg, 1.1x.1926, a.l., P. Tutein Nolthenius; 1 2, Nijmegen, 7.1x.1921, Lycklama a Nijeholt (ZMA); 1 d, Overveen, 29.viii.1930, G. A. Bentinck (RMNH).— Spain: 2 d, Andalucia, road to Istan, 400 m, 28.vi.1972, E. Traugott-Olsen; 1 9, idem, 200 m, 8.vii.1972; 2 &, Andalucia, road to Ca- sares, 500 m, 9.vii.1973, E. Traugott-Olsen (ETO). — Switzerland: 1 2, Erschmatt-Rotafen (Valais), 920 m, e:l. 7.vu.1983, mine Quercus pubescens 21.v.1983, S. E. Whitebread (coll. Whitebread). Mines.— Netherlands: Hollandse Radıng. Identity uncertain: Turkey: 1 9, Anatolia, Kizilca- hamam, 700 m, 31.vii-1.viti.1963, Arenberger (LNK) (specimen damaged). Additional records. — Italy: Latina, Monti Aurun- ci, Castelforte, 22.vi.1969, R. Johansson (adults at light); Piemonte (coll. Jackh) (R. Johansson, pers. comm.). 2. Ectoedemia (Zimmermannia) liebwerdella Zimmermann, 1940 (figs. 11, 36, 37 ,90, 145, 146, 232, 282, 330, 343, 348, 422, 473, 515) Ectoedemia liebwerdella Zimmermann, 1940: 264, 265, 1 fig. Holotype 2, Czechoslovakia: Deëín, (Tetschen) Liebwerd, 8.vi.1939, F. Zimmermann, Rindemine: Fagus silvat., Genitalia slide on pin (MHUB) [examined]. Ectoedemia liebwerdella; Klimesch, 1953: 195; 1961: 749; Schonherr, 1958: 1—71, figs. (detailed de- scription of all stages and biology); Lindner, 1959: 7—8 (Distribution in West-Germany); Szócs, 1965: 49; Haase, 1968: 61 (Distribution in East- Germany); Borkowski, 1975: 496. Zimmermannia liebwerdella; Hering, 1940: 266. Ectoedemia (Zimmermannia) lebwerdella; Hering, 1957: 437; Dorfmann, 1960: 17. Diagnosis: externally similar to longicaudella, but tornal and costal spots more distinct, expe- cially in female, and male with white hair-pen- cil. Differs from atrifrontella by brown thorax. Male genitalia extremely similar to atrifrontella, bud carinae hardly or not serrate, valva broader and ventral arms of transtilla longer. Female genitalia cannot be differentiated with certainty from atrifrontella. Description. Male (fig. 36). Forewing length 3.00—3.04 mm (3), wingspan 6.5—6.9 mm. Head: frontal tuft and collar dark brown to black. Antennae long, with 46—48 segments (2). Thorax dark brown to blackish fuscous. Forewings dark brown to blackish fuscous, almost uniform, sometimes slightly irrorate, tornal and to a less- er extent, costal spots white; cilia white beyond ill-defined cilia-line. Hindwing (fig. 11) with long white hair-pencil, of more than Ys hind- wing length, surrounded by white lamellar scales. Humeral lobe and costal emargination more pronounced than in other species. Female (fig. 37). Forewing length 3.60—3.64 mm (2), wingspan 7.8—8.4 mm. Antennal seg- ments 40—41 (2). Costal and tornal spot more pronounced than in male. Male genitalia (figs. 90, 330). Capsule length 377—429 um (4), slightly wider than in atrı- frontella: 291—343 um. Tegumen slightly cus- pidate. Gnathos (fig. 282) with central element long and narrow, parallel-sided. Valva (fig. 232) length 296—321 um (4), approximately triangu- lar, slender, but in comparison with atrifrontella wider, without any inner lobe, tip slightly hooked; transtillae with ventral arms interme- | diate in length between atrifrontella and longi- caudella. Aedeagus (figs. 343, 348) 454463 um (4), constricted at level of opening for duc- tus ejaculatorius; ventral carinae long, approxi- mately one third of total length aedeagus, with hardly serrate or smooth outer margins; lateral and dorsal carinae connected by prominent rim, stout and pointed, the dorsal longer. Vesica with distally a sclerotised plate with indistinct folds or ridges, in addition to small cornuti. Female genitalia (figs. 145, 146, 422). T8 with many long hairs, a row of 16—20 thicker and very long setae along anterior margin of T8, scales absent; anterior margin slightly indented. Anal papillae with 6—10 setae. Posterior apo- physes reaching slightly beyond anterior apo- physes. Vestibulum with pair of indistinct scle- rotisations. Corpus bursae + 1100 um, covered with pectinations, partly in concentric bands around signa; signa elongate, almost similar, length 390—495 um (6), 3.6—4.0 X as long as wide. Ductus spermathecae with 2'2—3 convo- lutions, becoming wider distally. Larva. Yellow, very elongate. Head-capsule brown. Ventral plates absent. See also Schonherr (1958). Biology. Host plant: Fagus sylvatica L. Mine (fig. 473). Contorted gallery in bark of trunks or thick branches. The larva feeds mainly in the direction of the main axis. Especially abundant on sunny side of trees. Life history. See excellent treatment by Schönherr (1958), larvae feed during two sum- Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 21 mers and overwinter twice to pupate in May- July, thus having a two-year cycle (in East Ger- many), but specimens completing their cycle in one year do occur (Schönherr, l.c). Adults emerge from early July to August. Distribution (fig. 515). Adults are only known from reared material from DDR, the holotype and French and Italian specimens collected at light. Records of mines known from East and West Germany, Silesia in Poland, Austria, Hungary, Italy: Alps and Apennines, France: Alps and Pyrenees. In northernmost Germany and Denmark the spe- cies could not be found, despite intensive search (Lindner, 1959; Schönherr, 1958). Material examined: 5 8,3 2. — Czechoslovakia: 1 2 Holotype, see above — France: 1 d, St. Barnabé, Col de Vence, 900 m (Alpes Marit.), 2—7.vu.1962, Arenberger (LNK). — Germany, East: 3 d, 2 9, Tharandt, el. 4—19.vii.1956, Fagus sylvatica, J. Schönherr (MHUB, 1 ZMC). — Italy: 1 &, Calabria- La Sila, prov. Cosenza, Longobucco, 1600 m, 3.v11.1982, at light, J. H. Kuchlein (coll. Kuchlein). Mines. — France: PEpine (Hautes Alpes); le Per- thus (Pyr.Or.); le Sappey-en-Chartreuse (Isere). Additional records. — Italy: Parco Nationale d’A- bruzzo, 1700—1800 m, mines, R. Johansson (pers. comm.); Trento, Mte Maranza, 10.x.1983, mines, E. J. van Nieukerken. 3. Ectoedemia (Zimmermannia) longicaudella Klimesch, 1953 19217, 21,27, 38,91, 147, 148, 233, 283, 331, 339,349, 34754235516) Ectoedemia longicaudella Klimesch, 1953; 193, 194, fig. 19. Lectotype d (here designated), Hungary: Nagy Nyir, Kecskemét, 17—28.v.1937, J. Kli- mesch, Genitalia slide Kl. 438 (ZSMK) [not exam- ined, genitalia figured by Klimesch]. Stigmella (Fomoria) peinii Nemes, 1972: 153—156, 1 fig. Holotype d, Rumania: Wald Gîrboavele, Bezirk Galati, 7.vii.1968, I. Nemes, Genitalia slide 1299 (coll. Nemes) [not examined]. Syn. nov. Trifurcula atrifrontella sensu auctt. partim. Ectoedemia longicaudella; Szócs, 1965: 50; Borkowski, 1970: 549, figs. 19, 26 (3 genitalia, externals); 1975: 496; van Nieukerken, 1982: 106, 107. Trifurcula (Ectoedemia) longicaudella; Johansson, 1971: 245. Ectoedemia (Zimmermannia) longicaudella; kowski, 1972: fig. 12 (venation). Bor- Diagnosis: the brown thorax and yellowish brown hair-pencil separate this species from atrifrontella, the hair-pencil and the absence of a costal spot from liebwerdella. From both spe- cies it is distinguished by the unconstricted ae- deagus, the shorter carinae, the wider capsule and longer ventral arms of transtilla in male, and by the long posterior apophyses and number of convolutions in spermathecal duct in female. See also hispanica and monemvasiae. Description. Male (fig. 38). Forewing length 2.68—3.64 mm (3.27 + 0.20, 28), wingspan 7.0—8.0 mm. Head: frontal tuft and collar dark brown to black. Antennae long, with 41—50 segments (45.2 + 2.5, 11). Thorax dark brown, often with white caudal tips of mesoscutum and tegulae. Forewings dark brown, irrorate with varying amount of white, tornal spot usually white; cilia silvery white beyond ill-defined cilia-line. Hindwing (fig. 12) with yellowish brown hair- pencil of approximately Vs hindwing length, surrounded by white lamellar scales; humeral lobe prominent, costal margin distinctly emar- ginated beyond hair-pencil Female. Forewing length 3.32—3.92 mm (3.67 + 0.19, 10), wingspan 7.2—8.6 mm. An- tennal segments 40—42 (41.2 + 0.8, 5). Male genitalia (figs. 91, 331). Capsule length 364—424 um (388 + 19.4, 15), wider than in atrifrontella, width 308—356 um. Vinculum with posterior part of ventral plate about half as long as ventral plate. Tegumen slightly cuspi- date. Gnathos (fig. 283) with central element long and narrow, parallel-sided. Valva (fig. 233) length 279—321 um (299.7 + 13.5, 15), triangu- lar, with indistinct rounded mesal lobe basally, not projecting beyond inner margin; transtilla with long ventral arm. Aedeagus (figs. 339, 340, 347) 343—403 um (435.7 + 19.1, 15), not con- stricted; ventral carinae long, but shorter than in atrifrontella, not serrate; lateral and dorsal carinae not connected by rim, stout and point- ed; dorsal carinae often bi- or multifurcate, with up to four horns each. Vesica with egg-shaped sclerotised plate in addition to small cornuti. Female genitalia (figs. 147, 148, 423). T8 with many long hairs, a row of more than 20 thicker and very long setae along anterior margin, scales absent; anterior margin of T8 almost straight, slightly indented. Anal papillae with 7—12 se- tae. Posterior apophyses reaching distinctly be- yond anterior apophyses. Vestibulum with pair of indistinct sclerotisations. Corpus bursae 1050—1450 um, covered with pectinations, partly in concentric bands around signa; signa elongate, similar, length 440— 737 um (562 + 77 22. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 um, 16), 4.0—5.2 x as long as wide. Ductus spermathecae with 34:—3% convolutions. Larva not examined. Biology. Host plants: Quercus robur L. and probably other Quercus species. Mines on Castanea could also belong to this species. In fact only reared once by Schönherr (1958), but mistaken for atrifrontella. “Mine. Not described, but probably not differ- ent from that of atrıfrontella. Life history. Under the name atrifrontella, Schönherr (1958) reported a two year cycle for this species — this is analogous to hiebwerdella. Adults are frequently collected at light in the months June and July, in Yugoslavia also in May, and occasionally in early August, thus not occurring as late as atrifrontella. Distribution (fig. 516). Widely distributed in central and southern Europe, but absent from Britain, and in Scandi- navia only known from southern Sweden. Not yet recorded from Portugal, Switzerland, Czechoslovakia, Bulgaria and Greece, but oc- curring in Anatolia. Remarks. This species was described from a long series covering many localities. Klimesch did not specify a holotype, but the identity of this spe- cies is clearly understood from his description and figure. Although I did not study the syn- types, I select here the specimen of which the genitalia were figured by Klimesch, as lecto- type, and therefore restrict the type locality to Nagy Nyir near Kecskemet, which also was listed as first locality in Klimesch’s list. Although I was not able to examine Stigmella (Fomoria) peruu Nemes, from the description and figure of male genitalia there is little doubt it isa synonym of longicaudella. Material examined: 40 46, 13 2, 1 ex. — Austria: 2 3, Gumpoldskirchen, Glaslauterriegel, 4.vu.1976, 18.vii.1980, F. Kasy; 8 d, Hackelsberg N. of Neu- siedlersee (near Jois), 23.vi.1975, 24.vi.1977 and 2.v1.1977, F. Kasy; 2 6, Hundsheimer Berg, Porta Hungarica (near Hainburg), 28.vi.1976, 2.vu.1977, F. Kasy (NMW). — Belgium: 1 2, Aye, 4.vu.1946, A. Richard; 2 2, Aye, 27.v11.1949, E. Janmoulle (IRSN). — lancer go Le “Nomina; (2 mear Diamo) 18.vii.1903, Chrétien; 3 d, Célé (Lot), 24—26 [= de- cades?], C. Dumont; 2 d, 1 2, Digne, vii—viii.1903, Chrétien snee MN 06 MR Ava land 16.vi.1917, Chrétien; 1 d, 2 ©, Revent. (interpreted as Reventin-Vaugris), 12—27.v11.1902, Chrétien (MNHN); 2 6, 1 2, St. Barrabé, Col de Vence (Alpes Marit.), 900 m, 2—7.v11.1962, Arenberger (ENIS) MES Viens Vaucluse) (nea pn), 10.viu.1974, R. Buvat (coll. Buvat). — Germany, East: 1 ©, Tharandt, el. 9.vii.1956, Quercus robur, J. Schönherr (MHUB). — Hungary, 1 d, Budakeszi, Härsbokorh., 24.vu.1952, L. Gozmany (MHUB); 1 36, Cserkut near Pécs, 12—20.vi.1936, J. Klimesch (LNK); 1 ex., Hu Nyírség, Bátorliget, 14.v1.1949, Kaszab & Székessy (MHUB); 1 dé, Kunadacs, 10.vi.1958, L. Kovacs (TMAB). — Netherlands: 4 6, Nijmegen, 14.vu.1926, 21.v11.1929, and 11.vu.1932. Lycklama a Nyeholt (RMNH, ZMA). — Spain: 4 d, 3 2, San Ildelfonso (La Granja), 8.vu.1902, Chrétien (MNHN). — . Sweden: 3 6, Högsby (Sm.), 17.vii.1976, R. Johansson (BMNH, EvN). — Turkey: 1 6, Anatolia, Kizilcahamam, 20.vi—8.vii.1970, Pink- r (LNK). — Yugoslavia: 1 2, Macedonia, Matka, Treschka Schlucht, 19—29.v.1955, J. Klimesch (ZSMK). Additional records. — Italy: Latina, Monti Aurun- ci, R. Johansson; Piemonte, Rocciamelone, 800 m, 8.vii.1961, at light, E. Jackh (both R. Johansson, pers. comm.). 4. Ectoedemia (Zimmermannia) hispanica sp. n. (figs. 39, 92, 234, 284, 332, 344, 345, 517) Type material: Holotype d, Spain: Andalu- cia, Sierra de Marbella, El Mirandor, 700 m, 14.vii.1980, E. Traugott-Olsen, Genitalia slide VU 1931 (ZMC). Paratype d, Spain: Aragon, Rubielos de Mora, 4.vii.1967, Arenberger (LNK). Diagnosis: male genitalia very characteristic with the pronounced lobe along inner margin of valva and broad and stout gnathos. Aedeagus similar to longicaudella. Externally character- ised by inconspicuous costal emargination and short hair-pencil. Description. Male (fig. 39). Forewing length 2.88—3.08 mm, wingspan 6.2—6.8 mm. Head: frontal tuft fuscous to dark brown. Antennae long, with 50—56 segments. Colour of thorax not unequi- vocal to determine (worn specimens). Fore- wings brown, probably uniformly coloured. Hindwing with relatively short white hair-pen- cil, about Y4 of hindwing length, surrounded by some white lamellar scales; humeral lobe less pronounced than in previous species, costal emargination very inconspicuous. Female unknown. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 23 Male genitalia (figs. 92, 332). Capsule length 334—374 um. Tegumen extended into rounded pseuduncus. Gnathos (fig. 284) with central el- ement wide and truncate. Valva (fig. 234) length 270—279 um, triangular, with prominent inner lobe in middle of valva; transtilla with long ven- tral arm. Aedeagus (figs. 344, 345) 377 um, slightly constricted in middle; ventral carinae long, as in longicaudella, not serrate; lateral and dorsal carinae not connected by rim, stout and pointed; dorsal carinae sometimes bifurcate. Larva unknown. Biology. Hostplant unknown. There are in the type- locality some large old Castanea trees and Quercus suber, but a search for barkmines in February 1984 was not succesful. Adults have been caught in July. Distribution (fig. 517). East and South Spain. Remarks. This species seems closely related to E. longı- caudella, but the male genitalia and hair-pencil are different enough to justify describing a new species. 5. Ectoedemia (Zimmermannia) monemvasiae sp. n. (figs. 13, 31, 32, 40, 93, 149, 150, 235, 285, 333, 349, 351, 352, 424, 538) Type material: Holotype 4, Greece (Hellas): Lakonia, 5 km s. Monemvasia, 28.v11.1979, G. Christensen, Genitalia slide VU 468 (ZMC). Paratypes: 5 6,4 9. — Greece: 1 2, Lakonia, 5 km s. Monemvasia, 1.viii.1978, G. Christensen; 2 ®, same data, but 28.vii.1979; 1 &, same data, but 8.vii.1979; 1 36, Lakonia, 7 km sw. Mo- nemvasia, 4.vii1.1979, G. Christensen; 1 9, same data, but 25.vii.1980; 1 d, same data, but 8.viu.1980 (ZMC, ZMA). — Turkey: 2 a, Anatolia, Kizilcahamam, 200 m, 31.vii— 1.v111.1963, Arenberger (LNK). Other material: Greece, 1 ? (abdomen and metathorax missing), Lakonia, 7 km sw Mo- nemvasia, 10.v11.1980, G. Christensen (ZMC). Diagnosis: male distinguished from the other Zimmermannia species treated here, by long brownish hair-pencil, surrounded by dark brown lamellar scales. Female by very dense bunch of long setae on abdominal tip dorsally. Male genitalia diagnosed by shape of vinculum, slender valvae with inner lobe, configuration of carinae and triangular cornutus and female geni- talia by number of convolutions in ductus sper- mathecae and hairy T8 and T9. Description. Male. Forewing length 2.84—3.36 mm (3.05 + 0.19, 5), wingspan 6.5—7.5 mm. Head: fron- tal tuft and collar fuscous. Antennae very long, with 49—58 segments (53.2 + 3.7, 5). Thorax and forewings brown, irrorate with white, sometimes an inconspicuous tornal spot white; cilia white beyond ill-defined cilia-line. Hind- wing (fig. 13) with long brown hair-pencil, al- most half as long as hindwing, surrounded by field of dark brown lamellar scales; humeral lobe prominent, costal margin with distinct emargination beyond hair-pencil. Female (fig. 40). Forewing length 2.6—3.0 mm (2.83 + 0.15, 5), wingspan 6.5—7 mm. An- tennal segments 42—44 (43.3 + 1.0, 4). Male genitalia (figs. 93, 333). Capsule length 386—429 um (3). Vinculum with ventral plate short, slightly excavate. Tegumen produced into blunt pseuduncus. Gnathos (fig. 285) with cen- tral element long and narrow, tapering towards sharp point. Valva (fig. 235) length 303—343 um (3), narrow triangular, with prominent inner lobe in middle of valva; transtilla with very long ventral arm. Aedeagus (figs. 349, 351, 352) 437—467 um (3), slightly constricted near opening of ductus ejaculatorius; ventral carinae long and parallel, fused near tip; lateral carinae small, almost triangular; dorsal carinae palmate, comprising each 4—5 teeth. Vesica with stout pointed triangular cornutus in addition to nu- merous small cornuti. Female genitalia (figs. 31, 149, 150, 424). Along anterior margin of T8 (? partly on T7) crescent shaped bundle of more than 50 very long setae, easily visible in undissected material, setae pectinate (fig. 32), on rest of T8 many short setae, scales absent. Anal papillae with more than 30 setae. Posterior apophyses reach- ing beyond anterior apophyses. Vestibulum wide, without distinct sclerotisations. Corpus bursae 1040—1080 um, covered with pectina- tions, partly in concentric bands around signa; signa elongate, slightly dissimilar, shortest 484—506 um, largest 583—616 um. Ductus spermathecae with 4/2—5 convolutions. Larva unknown. Biology. Hostplant: unknown, possibly a barkminer on Fagaceae. 24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Life history. Adults have been taken in July and early August. Distribution (fig. 538). Greece: Peloponnesos and Turkey: Anatolia. Remarks. This is a very distinctive species, of which several specimens of both sexes were collected from the type locality. The remarkably hairy abdominal tip of the female and the pectinate se- tae, suggest that this species lays its eggs on a very rough surface, such as old rugose bark. 6. Ectoedemia (Zimmermannia) amani Svens- son, 1966 (figs. 14, 41, 94, 151, 152, 236, 286, 334, 341, 342, 350, 425, 472, 474, 517) Ectoedemia amani Svensson, 1966: 200, 201, fig. 34, pl. 4 fig. 3. Holotype 4, Sweden: Sdm., Saltsjöba- den, 3.vii.1958, E. Aman, Genitalia slide 4107 (RMS) [examined]. Ectoedemia amani; Borkowski, 1975: 497, fig. 5 (d genitalia). Trifurcula (Ectoedemia) amanı; Johansson, 1971: 245. Trifurcula amani; Larsen, 1981: 71, 72, figs. 1—4 (4, 2 genitalia, distribution). Diagnosis: largest Ectoedemia from Europe, distinguished form preceding five species by orange head, absence of white spots on fore- wing and lower number of antennal segments. Differs from externally similar, but lighter, ligu- ricella, by presence of hair-pencil in male and lower number of antennal segments, in both sexes. Male genitalia characteristic with short and wide aedeagus, configuration of carinae and broad triangular valvae. Female genitalia espe- cially characterised by long spiraled ductus spermathecae, absence of long hairs on T8 and spines in vestibulum. Description. Male. Forewing length 3.2—3.92 mm (3.72 + 0.15, 6), wingspan 7.8—8.8 mm. Head: frontal tuft and collar orange to ochreous. Antennae not very long, with 36—41 segments (3). Tho- rax and forewing uniformly brown irrorate with white, without white spots; cilia lighter but ci- lia-line very inconspicuous. Hindwing (fig. 14) with snowwhite hair-pencil, approximately Y, of hindwing length, with a row of white scales along costal margin, but no specialised scales along dorsal edge; humeral lobe prominent, costal emargination present beyond hair-pencil. Female (fig. 41). Forewing length 3.84—4.52 mm (3), wingspan 8.8—9.8 mm. Antennal seg- ments 36—37 (3). Male genitalia (figs. 94, 334). Capsule length + 420 um (2), capsule very wide, 369—373 um (2). Vinculum with very short ventral plate. Te- gumen broadly rounded, not produced. Gnathos (fig. 286) with wide triangular central element. Valva (fig. 236) length 270—280 um (2), triangular, comparatively wide, tip curved slightly inwards, dorsal surface with indistinct serrate lobe. Aedeagus (figs. 341, 342, 350) 369—420 um (5), gradually widening from an- terior end towards wide posterior end; ventral carinae broadly triangular, separated, inner margin serrate; lateral carinae indistinct, round- ed; dorsal carinae comprising a row of 4—5 teeth; surface of aedeagus between ventral and lateral carinae with minute spines. Vesica with one broad triangular cornutus in addition to nu- ~ merous small cornuti. Female genitalia (figs. 151, 152, 425, 472). T8 with a row of 16—18 setae along anterior mar- gin and 4—10 small setae on disc, scales absent. Anal papillae with 15—21 seatae. Posterior apo- physes clearly reaching beyond anterior apo- physes. Vestibulum with two groups of spines, one near opening of ductus spermathecae and one opposite (fig. 472). Corpus bursae 1430— 1640 um, covered with pectinations, partly in concentric bands around signa; signa similar, 527—594 um (4) long, + 4 X as long as wide. Ductus spermathecae with 12'2—13 convolu- tions. Larva. Yellow, very elongate. Head-capsule brown. Ventral plates absent. Biology. Host plant: Ulmus spp. The species has not been reared, but often caught on Elm on which barkmines were observed (Johansson, pers. comm., Larsen, 1981). Mine (fig. 474). A long contorted gallery in smooth bark of rather thin branches, similar to that of atrifrontella. Life history. Not studied, but probably simi- lar to that of Lebwerdella. Adults have been caught in June (southern Europe only) and July. Distribution (fig. 517). Recorded from southern Norway (see below, not on map), southern Sweden, Denmark: Bornholm and Falster, Austria: Vienna region, and Yugoslavia: Macedonia. | Kritzendorfer Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 25 Remarks. Although one of the largest nepticulid spe- | cies, E. amani was only discovered in 1966 by | Svensson in Sweden. Since then several speci- mens have been found in Sweden and Denmark (Larsen, 1981). Outside Scandinavia only the four specimens cited below are at present known, plus the larva and mines found near Bad Deutsch Altenburg. This species resembles E. piperella Wilkinson & Newton, 1981 from USA. Material examined: 8 6, 4 2. — Austria: 1 6, Hundsheimer Berg, Porta Hungarica (near Hain- burg), 23.vii.1977, F. Kasy; 1 dg, Klosterneuburg, Au, 29.v1.1936, al, Preissecker (NMW). — Sweden: 1 & Holotype, see above; 2 d, 2 ©, Kullaberg (Sk.), 19.vi1.1974, 11—12.v11.1975, R. Johansson (BMNH, EvN); 2 6, Stockholm, Skogskyrkogd, 4.vi1.1973, B. Gustafsson; 1 6, Upland, Riksmuseet, 10.vii.1973, B. Gustafsson (RMS). — Yugoslavia: 1 2, Macedonia, Stari Dojran, 10—19.vi.1955, J. Klimesch (ZSMK); 1 2, Macedo- nia, Treschka Schlucht near Skopje, 1—8.vii, F. Kasy (NMW). Larva and mines. — Austria; 1 final instar larva, mines, Bad Deutsch Altenburg, W. of Hainburg, Pfaf- fenberg, 23.x.1983, E. J. van Nieukerken (ZMA). Additional record. — Norway; Ak., Baerum, Ostoya, 1 d, 2—9.v11.1983 (Johansson, in litt.). 7. Ectoedemia (Zimmermannia) nuristanica sp. n. (figs. 42, 95, 153, 154, 237, 287, 335, 353—355, 426) Type material: Holotype d, Afghanistan: Nuristan, 25 km N. Barikot, 1800 m, 12— 17.vi1.1963, Kasy & Vartian, Genitalia slide MV 5402 (NMW). Paratype ®, same data (NMW). Diagnosis: the only known dark-headed (Pal- aearctic) Zimmermannia without hair-pencil in | male. Male genitalia characterised by pointed pseuduncus, narrow valvae and three pairs of al- most similar carinae. Female characterised by very dense bundle of extremely long setae on tergite 7. Description. Male holotype (fig. 42). Forewing length 2.84 mm, wingspan 6.4 mm. Head: frontal tuft and collar dark brown. Antennae broken. Thorax and forewings brown irrorate with white, with an inconspicuous white dorsal spot. Hindwing without hair-pencil, costal bristles or specialised scales; humeral lobe more or less distinct. Female paratype. Forewing length 3.08 mm, wingspan 7 mm. Antennae long, with 41 seg- ments. Male genitalia (figs. 95, 335). Capsule length 403 um, width 261 um. Tegumen produced into cuspidate pseuduncus. Gnathos (fig. 287) with long, slender central element (in figure not in proper ventral view). Valva (fig. 237) length 266 um, narrow triangular, with indistinct inner lobe (mesal), distally suddenly narrowed into fingerlike tip. Aedeagus (figs. 353—355) 351 um, hardly constricted; ventral carinae short, widely separate, bifurcate; lateral and dorsal ca- rinae similar in size and shape, horn-shaped, closely placed. Vesica difficult to study in holo- type, no special cornuti visible. Female genitalia (figs. 153, 154, 426). T7 with horseshoe-shaped dense bundle of extremely long setae, reaching beyond abdominal tip. T8 with a row of about 20 long setae along anterior margin and with many shorter setae on disc. Anal papillae with 30—32 setae. Posterior apo- physes hardly reaching beyond anterior apo- physes. Vestibulum with indistinct sclerotisa- tion. Corpus bursae 935 um long, covered with pectinations, partly in concentric bands around signa; signa similar, 399 and 424 um long, 4.5— 4.65 X as long as wide. Ductus spermathecae with 412 convolutions. Larva unknown. Biology. Hostplant: unknown. The specimens were taken at light in mountains with extensive woods of Quercus baloot Griff., a relative of Q. ilex L. (Kasy, 1965), it is therefore possible that nuristanica is a barkminer of Q. baloot. Life history. Adults taken in July. Distribution. Only known from East Afghanistan: Nuris- tan. Remarks. It is assumed that both sexes described here belong to the same species, since they are exter- nally similar and have been collected together. 8. Ectoedemia (Zimmermannia) liguricella Klimesch, 1953 (figs. 43, 96, 155, 156, 238, 288, 336, 356—358, 427, 428, 539) Ectoedemia liguricella Klimesch, 1953: 194, 195, figs. 20—22. Lectotype d (here designated), Italy: Li- guria, prov. Savona, Noli, v or 1x.1951, J. Kli- 26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 mesch, Genitalia slide Kl. 513 (ZSMK) [not exam- ined, genitalia figured by Klimesch]. Ectoedemia liguricella; Sz6es, 1965: 49. Diagnosis: differs from all treated Zimmer- mannia species, except amani, by light coloured head. Males can be separated from amanı by ab- sence of hair-pencil, and females by larger num- ber of antennal segments. Diagnostic in male genitalia are the vinculum process, the short narrow gnathos, the shape of the valva and the configuration of the carinae. The female genita- lia are characterised by the relatively few setae on T8 and the 4/—512 convolutions of the ductus spermathecae. E. liguricella can be con- fused with Trifurcula species, see generic diag- nosis. Description. Male (fig. 43). Forewing length 3.0—4.04 mm (3.58 + 0.29, 12), wingspan 7.6—8.8 mm. Head: frontal tuft and collar yellow to yellow ochreous. Antennae long, with 43—48 seg- ments (44.9 + 1.7, 7). Thorax and forewings brown irrorate with yellowish-white (European specimens darker than Moroccan), with sometimes small indistinct white tornal spot; cı- lia-line hardly visible. Hindwing without hair- pencil, costal bristles or special scales. Humeral lobe distinct, rounded. Female. Forewing length 3.44—4.0 mm (3.73 + 0.24, 6), wingspan 7.6—9 mm. Antennal seg- ments 39—44 (41.6 + 2.9, 7). Male genitalia (figs. 96, 336). Capsule length 321—377 um (357 + 20.4, 10). Vinculum with ventral plate narrow. Tegumen rounded, with an obvious anteriorly directed, tongue-shaped process. Gnathos (fig. 288) with narrow pointed central element, shorter than in related species. Valva (fig. 238) length 270—304 um (284.8 + 12.7, 9), narrow triangular, with distinct inner (mesal) lobe in middle; transtillae with short transverse bar. Aedeagus (figs. 356—358) 369— 420 um (398.1 + 16.7, 10), slightly constricted; ventral carinae long, widely separate, pointing outwards; lateral carinae absent; dorsal carinae simple, pointed; aedeagus dorsally ending in two weakly sclerotised lobes covered with spin- es, less spines on left lobe. Vesica with small cornuti only. Female genitalia (figs. 155, 156, 427, 428). T8 with a row of about 10—20 relatively long se- tae, along anterior margin, and with a row of 10—20 shorter setae more posteriorly, scales absent. Anal papillae with 15—27 setae. Posteri- or apophyses reaching beyond anterior apo- physes. Vestibulum with indistinct internal scle- rotisation. Corpus bursae 880—1100 um, cov- ered with pectinations, especially dense in ductus bursae, partly in concentric bands around signa; signa similar, 308—493 um (395.9 + 59.4, 8), 5.57.0 x as long as wide. Ductus spermathecae with 4/2—512 convolutions. Larva unknown. Biology. Hostplant: unknown. It might be a barkmin- er of evergreen Quercus, since it has often been collected amongst those trees. In one of the lo- calities near Marbella I noted a few barkmines on Quercus coccifera, which could belong to E. liguricella. Life history. Adults taken from May to Sep- tember. Distribution (fig. 539). A western mediterranean species, known from the Italian Riviera, France, Spain and Mo- rocco. Occurs from sea-level to high elevations in the mountains (1600 m in Spain, 2600 m in Morocco). Remarks. I have not examined any types of liguricella, because the identity of this species is clear from Klimesch’s (1953) figure of the male genitalia, and hence, the specimen represented by that fig- ure is here selected as lectotype. For the first time the species is here recorded from areas out- side the type-locality. The female collected in the company of 6 males in Morocco has slightly different genitalia (fig. 428) from the Spanish specimens, and is therefore not included in the measurements of the female genitalia. It has 80 setae along the an- terior margin of T8, 45 setae more posterior on T8 and anal papillae with 39 setae. The bursa is smaller, 715 um, with signa of 283 and 317 um. The total appearance of the specimen however, does not indicate that it is a different species, but more material is needed to see if this varia- tion is constant. Material examined: 39 d, 21 9. — France: 2 dg, “Nesp.” (? near St. Pons, dep. Hérault), 15.vi.1904, Chrétien; 1 6, Ile du Levant (Var), 19.vu.1941, H. Legrand (MNHN). — Morocco: 6 d, 1 2, Haut Atlas, Oukaim’den (near Toubqual), 2600 m, 9— 11.vu.1975, F. Kasy (NMW). — Spain: 1 d, Albarra- cin, Noguera, 1600 m, 18—22.vii.1960, Vartan (NMW); 3 6, Andalucia, road to Benahavis, VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 27 8.v1.1983, E. Traugott-Olsen; 1 9, Andalucia, road to Casares, 500 m, 9.vii.1973, E. Traugott-Olsen; 2 d, 4 2, Andalucia (Marbella-region), road to Istan, 400 m, data: 17.vii.1971, 21.v1.1972, 4.v1.1973, 25.vi.1975, 15.vii.1982, E. Traugott-Olsen; 2 d, 6 9, Andalucia (Marbella-region), road to Ojen, 150 m, data: 5.v.1980, 12 and 25.vi.1981, 20 and 25.v1.1983, E. Traugott-Olsen; 1 9, Andalucia (Marbella-region), Refugio de Juanar, 700 m, 29.vii.1971, E. Traugott- Olsen, 2 &, 1 2, Andalucia, Sierra de Marbella, El Mirandor, 700 m, 21.vii.1982, E. Traugott-Olsen (ETO, ZMA, ZMC, EvN); 1 6, Andalucia (Gra- nada), Sierra de Alfacar, 1200 m, 26.vi—8.vii.1962, W. Glaser (LNK); 1 ©, idem, 1500 m, 23.vi.1968, K. Sattler & D. J. Carter (BMNH); 1 6, Aragon, Ru- bielos de Mora, 4.v11.1967, Arenberger; 2 d, 5 &, Cataluna, Port Bou, 11—18.vii.1967, Arenberger; 3 6, idem, 0—300 m, 9—24.vi.1964, M. & W. Glaser (LNK); 1 6, Huelva prov., Torre la Higuera, 12.v.1981, C. Gielis (coll. Gielis); 12 d, 1 2, idem, 221v—9.v.1983. J. B. Wolschrijn (coll. Wolschrijn, ZMA, EvN). Subgenus Ectoedemia Busck Ectoedemia Busck, 1907: 97. Type-species: Ectoede- mia populella Busck, 1907: 98; by original desig- nation and monotypy. Dechtiria Beirne, 1945: 204. Type-species: Tinea sub- bimaculella Haworth, 1828: 583; by original des- ignation. (Synonymised by Svensson, 1966: 200). Ectoedemia (Dechtiria); Borkowski, 1972: 699; Em- met, 1976: 188, 191. Ectoedemia (Ectoedemia); Borkowski, 1972: 699; Emmet, 1976: 188, 189; Scoble, 1983: 20. Ectoedemia; Scoble, 1978: 82; 1979: 35—54; Wilkin- son & Scoble, 1979: 73; Wilkinson & Newton, 1981: 32 partım. Trifurcula (Ectoedemia); Johansson, 1971: 245. Description. Adult. Small to moderately large nepticulid moths, forewing length 1.7—3.7 mm (wingspan 3.28.4 mm). Head. Antennae short or long; in male with 24—63 segments, in female with 21—43. Wings. Colour pattern variable, often a white medial fascia or costal and dorsal spots present, sometimes basal or discal spot in addition, sometimes white markings absent. Cilia-line present except in occultella-group and populella. Hindwing in male either with costal bristles or hair-pencil, in some species both absent. Addi- tional special scales occur in several species. Humeral lobe not very prominent, or absent. Forewing venation (fig. 9). R and M + Cu forming closed cell, branches R,, R,,3, Ry, Rs, M and Cu present. A thickened, without anal loop. Cu and A in some species very long, seeming fused at tips. Male genitalia. Vinculum ring shaped, ante- rior extension short, anteriorly convex. Tegu- men produced into distinct pseuduncus, of vari- able form. Uncus absent. Gnathos with spatu- late or triangular central element, sometimes divided into a distal spatulate part and basal part with serrate margins. Valva approximately tri- angular, or almost rectangular, with tip directed inwards or posteriorly, often clearly separate from rest of valva. No mesal (inner) lobes pre- sent. Aedeagus in all but one species with ven- tral carinae, often bi- or multifurcate, and in some species in addition with dorsolateral cari- nae. Vesica in all but one species with numerous small denticulate cornuti only: Female genitalia. Tergite 7 with or without a row of long setae near anterior margin of tergite 8. Tergite 8 often with two patches of setae and scales, sometimes with setae only. Anal papillae with setae. Vestibulum in most species with ring-shaped vaginal sclerite and denticulate pouch. Corpus bursae with numerous pectina- tions, or pectinations concentrated posteriorly near vestibulum. Reticulate signa present, of variable form and often dissimilar. Margin of signa narrower than individual cells. Ductus spermathecae spiraled, with variable number of convolutions. Larva. Yellow, white, green or grey, feeds venter upwards. Probably all species have four larval instars. Many species have 12 sclerotised ventral plates during second and third instar, being shed in the final instar, independently from moult. In some species similar dorsal plates occur in addition. Biology. Larva leaf-miner, or petiole-miner. Western Palaearctic species mine on Fagaceae, Rosaceae, Salicaceae and to a lesser extent on Betulaceae, Ulmaceae and Anacardiaceae. In addition spe- cies from other regions are recorded from Nys- saceae, Platanaceae, Juglandaceae, Aceraceae, Hippocastanaceae, Ericaceae, Caprifoliaceae and Burseraceae. Most European species are univoltine, feeding in late summer or autumn, but al least terebinthivora is bivoltine, and some others are suspected to be so. Larvae of many species are often gregarious. Larvae overwinter full-fed in cocoons in the soil, or in the mine in the case of agrimoniae and pupate in spring. Adults emerge in March—July. Some mediter- ranean species, on evergreen oaks, feed in the winter and aestivate in their cocoons, or emerge in spring. 28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Distribution and composition. The distribution is mainly Holarctic: 42 spe- cies are reported here from the West Palaearc- tic, Wilkinson & Newton (1981) and Wilkinson (1981) reported 18 North American species, Pu- plesis (1984a and b) described 9 species from the eastern USSR and about 25 species occur in a collection of Japanese Nepticulidae. In addition three species are known from Southern Africa (Scoble, 1978; 1979). The Ectoedemia populella group This group comprises all the Salicaceae-feed- ing Ectoedemia species. Most make mines in the petiole and later in the lamina of Populus spe- cies, intimella makes a similar mine on Salix but starts in the midrib, not the petiole, and the Nearctic populella makes a petiole-gall. All feed late in the year and are often found in the green islands of fallen leaves. Adults are often found resting on trunks. Male genitalia are characterised by the pres- ence of two pairs of carinae, which are often large. Female genitalia are characterised by the presence of a vaginal sclerite, a spiculate pouch with conspicuous and equally spaced spicules and a bursa, usually covered with pectinations (except intimella). The ductus spermathecae has 212—3 convolutions and the signa are elongate and almost similar. Males invariably possess a hair-pencil on the hindwing. The group is Holarctic, and comprises also the Nearctic E. populella Busck and E. canutus Wilkinson & Scoble and the Eastern Palaearctic E. wilkinsoni Puplesis, 1984a. Ectoedemia (Ectoedemia) populella Busck, (figs. 98, 240, 360) Ectoedemia populella Busck, 1907: 98. Ectoedemia populella; Borkowski, 1972: 697; Wilkin- son & Scoble, 1979: 74—77, figs. 41, 42; Wilkin- son & Newton, 1981: 41, figs. 4, 5. E. populella does not occur in the Western Palaearctic Region, but is treated here because it is the type-species of Ectoedemia. A full de- scription is given by Wilkinson & Scoble (1979). Some descriptive notes are given in order to compare it with the Western Palaearctic species. Adult. Antennae very long, with approxi- mately 63 segments in d and 42—43 in 2. Fore- wings including cilia uniform cupreous brown, no cilia-line, hindwing in d with short incon- spicuous brown hair-pencil. Male genitalia (figs. 98, 240, 360). Capsule length + 390 um. Tegumen produced into rounded pseuduncus. Gnathos with smooth spatulate, slightly truncate central element. Val- va (fig. 240) length + 215 um, broad, tip hardly demarcated, with many setae; inner margin slightly sinuous. Aedeagus (fig. 360) + 380 um, with long pointed ventral carinae and very simi- lar dorsolateral carinae. Female genitalia. Terminalia very wide. T7 without row of setae. T8 wide, with two patches of scales and 10 setae at least. Anal pa- pillae with 7—11 setae. Vestibulum with vaginal sclerite, a spiculate pouch with many short, sin- gle denticles and a dense patch of pectinations near entrance of ductus spermathecae. Corpus bursae without pectinations; signa comparative- ly short, + 270—320 um, cells very spiny. Duc- | tus spermathecae broken in single slide exam- ined. Remarks. E. populella makes petiole-galls in several Populus-species. In some characters it is aber- rant in comparison with European species such as absence of cilia-line, large number of anten- nal segments. Material examined. — USA: 4 6, 2 9, syntypes, no. 3238, 12—24.v.1884, Poplar (USNM). 9. Ectoedemia (Ectoedemia) intimella (Zeller, 1848) (es 09/4407 1571585 2395 289 SSP ONE 520) Nepticula intimella Zeller, 1848: 323. Holotype ©. Poland: Glogöw (Glogau), Zeller (depository un- known) [not examined]. Nepticula intimella; Stainton, 1849: 29; 1854: 299; Herrich-Schaffer, 1855: 356; Frey, 1857: 393, 394: Stainton, 1859: 432; Wocke, 1871: 339; 1874: 102; Nolcken, 1871: 792; Heinemann & Wocke, 1877: 764; Sorhagen, 1886: 309; Meyrick, 1895: 724, 725; Tutt, 1899: 341, 342; Rebel, 1901: 227; Meess, 1910: 480; Sorhagen, 1922: 54, pl. 3 fig. 61; Meyrick, 1928: 861; Petersen, 1930: 74, fig. 110 (d genitalia); Hering, 1943: 275, fig. 2 (d genitalia); Szöcs, 1965: 82. Dechtiria intimella; Beirne, 1945: 205, fig. 67 (6 geni talia); Emmet, 1971: 280, 281. Stigmella intimella; Klimesch, 1951: 63, 64; Gerasi- mov, 1952: 244; Klimesch, 1961: 762; Lhomme, 1963: 1199; Borkowski, 1969: 112. Stigmella (Dechtiria) intimella; Hering, 1957: 811, 928, fig. 588b (mine). Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 29 Trifurcula (Ectoedemia) intimella; Johansson, 1971: 245. Ectoedemia intimella; Bradley et al., 1972: 3; Borkowski, 1975: 494; Emmet, 1976: 190, pl. 7 fig. 1, pl. 12 fig. 34; van Nieukerken, 1982: 107. Trifurcula intimella; Karsholt & Nielsen, 1976: 18. Diagnosis: easily separated from most Ectoe- demia species by presence of a medial dorsal spot only on forewing. Distinguished from ilicis and heringella by more uniformly dark fore- wings, hair-pencil in male, and the flagellum be- ing the same colour as the scape: it is the only treated Ectoedemia, with this character. Species of Fomoria, Stigmella or Ectoedemia (Zimmer- mannia) with dorsal spot only, have it in post- medial position. Description. Male. Forewing length 2.4—2.84 mm (2.58 + 0.15, 16), wingspan 5.3—6.3 mm. Head: frontal tuft and collar intensively ferruginous to yel- | lowish orange. Antenna with 39—45 segments . (41.3 + 2.0, 12), scape, pedicel and flagellum | yellowish white, with an orange tinge. Thorax and forewings uniformly blackish fuscous, with a faint purplish gloss, scales almost uniformly dark; a yellowish white dorsal spot in middle of forewing, conspicuous. Hindwing with a very short ochreous hair-pencil, less than 1/5 of hindwing length. Female (fig. 44). Forewing length 2.48—3.04 mm (2.75 + 0.19, 7), wingspan 5.6—6.8 mm. Antenna with 27—30 segments (28 + 1.1, 6). Ovipositor protruding, pointed. Male genitalia (figs. 97, 239, 289, 359). Cap- sule length 287—304 um (294.3 + 8.0, 6). Tegu- men produced into wide, triangular pseudun- cus. Gnathos (fig. 289) with central element very wide, uniformly rounded. Valva (fig. 239) length 210—236 um (217.1 + 9.3, 6), basally broad, suddenly narrowed in middle with inner margin becoming strongly concave; tip pointed. Aedeagus (fig. 359) 317—364 um (340.3 + 18.3, 5), with pair of slender, pointed ventral carinae, sometimes bifid, and pair of pointed dorsolat- eral carinae with additional spines. Female genitalia (figs. 157, 158, 429). T7 with a row of 6—8 setae along posterior margin. T8 narrowed posteriorly, with two lateral groups of 11—16 short and long setae. Anal papillae narrow, with 14—15 setae. Vestibulum with va- ginal sclerite, a dorsal spiculate pouch with comparatively few (less than 40) spines, all sin- gle and equally spaced; patch of densely packed pectinations near opening of ductus spermathe- cae. Corpus bursae 505—605 um, without pec- tinations; signa dissimilar, longest 304— 347 um (4), shortest 257—313 um (4), 4.9—5.5 X as long as wide. Ductus spermathecae with 212—3 convolutions. Larva. Pale yellow. Sternites present on pro- and mesothorax and abdominal segment 10. Ventral plates absent. Biology. Hostplants. Salix caprea L., S. cinerea L., S. pentandra L., S. fragilis L., and S. phylicifolia L Mine (fig. 477). Egg on upperside, against midrib. Early mine in midrib, later becoming large elongate blotch at one side of midrib, with black frass deposited in two lateral lines, such that larva can pass in between to conceal itself in midrib. Only final instar larva mines in leaf- blade. Life history. Univoltine. Larvae feed late in the season, from late September until Novem- ber, often in green islands in fallen leaves. Adults in June and July. Distribution (fig. 520). Widely distributed in northern, western and central Europe, but not yet recorded from Nor- way and Ireland. In the south only known from northern Italy, North Yugoslavia and Rumania. Remarks. There is unfortunately no specimen in the Zeller collection in BMNH, which can be re- garded as the holotype. Zeller’s description 1s however very clear, since he amongst others noted the completely yellow antennae, which are very characteristic for intimella. Conse- quently the identity of this species has never been in doubt. Material examined: 31 d, 23 2, 1 ex. — Austria: 1 3, Hirschdorf, Ob. Ost., el. 11.v. 1898, Hauder; 1 9, Klosterneuburg, Freiberg, e.l. 28.v.1941, Salix caprea, Preissecker (NMW). — Germany, East: 2 d, 1 2, Berlin, Finkenkrug, el. 27.ii—7.111.1918, Salix caprea, Hering; 1 dg, 1 ©, Bredow near Nauen, el. 31.v— 2.vi.1923, Hering (MHUB); 1 &, Görlitz, 24.vi.1884 (NMW); 1 8, Rachlau, Schütze (ZSM); 2 6, 3 ®, Rachlau, 1902, Salıx caprea, Schütze (MHUB). — Great Britain: 2 6,1 2, 2 km SE Earls Colne: Chalk- ney Wood (Essex), e.l. 30.v—16.vi.1980, Salix caprea, Bryan, Emmet & van Nieukerken (ZMA). — Nether- lands: 1 ©, Amsterdamse Bos, e.l. 24.v1.1983, Salix cinerea, J. Brouwer; 1 d, 2 2, Ootmarsum: Ageler- 30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 broek, e.l. 8—12.v.1982, Salix cinerea, Andeweg & van Nieukerken; 4 &, Rockanje: Voornes Duin, e.l. 17.vi—4.vii.1980, Salix cinerea, van Nieukerken; 3 d, Schinveld, 29.vi.1975, G.R. Langohr (ZMA); 1 2, Zwanewater, 5.vu.1982, Koster (coll. Koster). — Po- land: 8 6, 5 ©, 1 ex, Wroclaw (Breslau), el. 11 — iv.1875, Salix fragilis, Wocke (MHUB, NMW, RMNH, ZSM); 1 6,1 ©, Silesia (MHUB). — Swit- zerland: 1 ©, St. Gallen, e.l. iv.1915, Müller-Rutz (ZSM). — Yugoslavia: 2 d, 3 9, Mt. Slavnik, 8 km S. Herpelje-Kozina (Slovenia), + 900 m, e.l. 10— 15.vi.1984, Salix caprea, J.J. Boomsma & E. J. van Nieukerken (ZMA). Mines. On Salix caprea: Austria: Nassfeld Pass, SW Hermagor.— Belgium: Zolder. — Great Britain: SE Earls Colne. — Yugoslavia: Mt. Slavnik, S. Herpelje- Kozina. On Salix cinerea: Netherlands: Aalsmeer; Amsterdamse Bos; Ootmarsum; Rockanje. 10. Ectoedemia (Ectoedemia) hannoverella (Glitz, 1872) (ES, Je AAR AS, OD, USD 160 ZAAL MIO soils SY) 430, 475, 518) Nepticula hannoverella Glitz, 1872: 25, 26. Lectotype d (here designated), Germany: Hannover, Glitz, coll. Staudinger, Genitalia slide 1521 RJ (MHUB) [examined] Nepticula hannoverella; Wocke, 1871: 340; 1874: 103; Heinemann & Wocke, 1877: 766; Rebel, 1901: 227; Meess, 1910: 480; Sorhagen, 1922: 58; Petersen, 1930: 76, fig. 116 (d genitalia); Hering, 1935: 7; Szöcs, 1965: 85. Stigmella hannoverella; Klimesch, 1951: 64; Gerasi- mov, 1952: 241; Klimesch, 1961: 763; Lhomme, 1963: 102; Borkowski, 1969: 107. Stigmella (Dechtiria) hannoverella; Hering, 1957: 811 (mine). Trifurcula (Ectoedemia) hannoverella; Johansson, 1971: 245. Ectoedemia hannoverella; Borkowski, 1972: fig. 7 (6 genitalia); 1975: 495; van Nieukerken, 1982: 107, figs. 1,5 (5 genitalia, mine). Diagnosis: externally easy to confuse with turbidella, but in female the blunt ovipositor of hannoverella separates it immediately from tur- bidella, which has a pointed ovipositor. Males with dark heads always belong to turbidella, but light-headed males can only be separated by the genitalia. These are very different in shape of valva, shape and size of carinate processes, and gnathos, which bears spines in hannoverella. From other Ectoedemia species hannoverella and turbidella can be separated by the presence of a white discal spot in basal part of forewing and many scattered white scales; males also pos- sess a hair-pencil. Description. Male (fig. 45). Forewing length 2.4—3.16 mm (2.84 + 0.21, 18), wingspan 5.2—6.8 mm. Head: frontal tuft yellowish orange to light fer- ruginous; collar slightly lighter. Antennae with 4453 segments (48.8 + 2.8, 10). Thorax fus- cous black with some white scales along frontal margin; forewings fuscous black with a variable pattern of yellowish white spots; usually a me- dial costal and opposite dorsal spot, sometimes fused by some, more distally placed, scales; bas- al half with many scattered white scales, often forming a small discal spot halfway between wingbase and costal spot, and a basal spot along dorsal margin. Specimens with almost uniform dark forewings occur. Hindwing with a yellow- ish-white hair-pencil, about 1/5th of hindwing length. Female. Forewing length 2.8—3.32 mm (3.05 + 0.16, 14), wingspan 6.2—7.2 mm. Antennae’ with 29—33 segments (30.9 + 1.6, 9). Male genitalia (figs. 99, 241, 290, 361, 399). Capsule length 249—309 um (282.9 + 20.8, 6). Tegumen wide and rounded. Gnathos (fig. 290) with moderately long central element, ventrally with some rows of spines. Valva (fig. 241) length 201—236 um (217.3 + 11.8, 7), inner margin almost straight, except basally; outer margin strongly convex, widest part beyond middle; apex of valva not separated, hardly curved inwards, forming an almost right angle. Aedeagus (figs. 361, 299) 291—339 um (309.8 + 16.2, 7), with two pairs almost similar pointed carinae, hardly curved, without additional spines. Female genitalia (figs. 159, 160, 430). 17 without row of setae. T8 broad, rectangular or trapezoid, with two lateral patches of scales and at least 12—17 setae. Anal papillae with 6—11 setae. Vestibulum strongly folded stained by chlorazol), with vaginal sclerite, dor- sal spiculate pouch with + 50 single and equally spaced spines, and a patch of densely packed pectinations near entrance of ductus spermathe- cae. Corpus bursae long and slender, 660—880 um, covered with pectinations, partly in con- centric bands around signa, absent in anterior part; signa almost similar, 390—480 um (422.1 + 26.4, 10), 3.43.7 X as long as wide. Ductus spermathecae with 212—3 convolutions. Larva. Pale yellow. All thoracic segments and abdominal segments 8—10 with light brown sternites. Ventral plates absent. (heavily | Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 31 Biology. Hostplants: Populus nigra L. and its hybrids (P. x canadensis Moench.) Mine (fig. 475). Egg deposited on lateral side of petiole, about one centimeter from lamina. Mine first straight gallery in petiole, causing swelling; in final instar larva enters lamina, making elongate blotch, usually between first lateral vein and leaf margin, occasionally be- tween midrib and first lateral vein; frass in two | parallel lines, leaving passage for larva, which can withdraw itself in petiole. Live history. Univoltine. Larvae start feeding early, probably already in July, but feed very | slowly; blotches with final instar larvae can be found from late September to November, often in green islands of fallen leaves. Larvae feed | usually in the dark. Adults in May and June. Distribution (fig. 518). Only known from a comparatively small area in central Europe, where it is widespread and often abundant. Absent from the British Isles and Scandinavia, but known from Denmark. Only two records from France, and not yet re- corded south of the Po valley in Italy or south of the Danube in Yugoslavia. Buszko (in litt.) suggests that the species is expanding its area, on the basis of an increase of records in Poland. Remarks. Since types no longer exist in the Glitz collec- tion in Hannover (Niedersächsisches Landes- museum), a lectotype is selected from specimens in the Staudinger collection. Material examined: 66 6,56 2. — Austria: 2 6, 1 9, Klosterneuburg, Kritzendorfer Au, el. 11— 15.v.1937, 24.iv.1938, Preissecker (NMW); 1 4, Wien, Mann, Zeller coll. (sub turbidella) (BMNH); 1 2, Wien, Prater, el. 10—12.v1.1984, E. J. van Nieu- kerken (ZMA). — Belgium: 1 2, Elewijt, 20.v1.1944, | L. Legiest; 1 d, Laeken (Brussel), 9.v.1945, L. Legiest (IRSN); 2 d, 1 2, S. of Rouvreux (Liège), el. 26— | 27.v.1980, Bryan & van Nieukerken (ZMA). — france: 1 8, 1 2, Alpes Maritimes, Toet s.Var, 10.v.1980, C. Gielis (coll. Gielis). — Germany, West: 1 g, Baiern, 1858 (NMW); 1 6, 1 ©, Grünnstadt, Pfalz, Eppelsheim (ZSM); 1 d, Hannover, Heine- | mann (RMNH); 1 ©, Hannover, (NMW); 2 d, 1 2 (lecto- and paralectotypes), Hannover, Glitz, coll. Staudinger (MHUB); 2 6, 2 2, Regensburg, D. O. Hofmann (RMNH); 3 d, Regensburg, Frank (ZSM); 1 ©, Regensburg, 28.v.1885 (NMW); 6 d, 3 ©, Re- gensburg (MHUB). — Germany, East: 1 ©, Bautzen, 2.11.1907 (NMW); 4 6, 9 2, Berlin-Dahlem, el. 28.11— 10.11.1958, Hering; 5 d, 8 9, Bredow near Nauen, el. 22.11—16.1v.1924, 1.vu.1923, Hering (MHUB); 1 2, Erfurt, e.l. 1884 (RMNH). — Nether- lands: 19 6, 15 2, from following localities: Amster- damse Bos; Bunde; Geulle; De Lutte; Oostvoorne; Susteren; Winterswijk; Zwanewater (ZMA, coll. Kos- ter). — Poland: 4 6, 1 ©, Wroclaw (Breslau), e.l. 11.1868, 11.1869, [Wocke] (MHUB, ZMA). — Swit- zerland: 1 à, Landquart, e.l. 26.1v.1916, Müller-Rutz (ZSM). — Yugoslavia: 2 9, 2 km W of Bezdan (Voj- vodina), valley of Danube, el. 16—19.v1.1984, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. — Austria: Klosterneuburg; Mühlleiten (Grossenzersdorf). — France: Schirmeck. — Ger- many, West: Hillesheim. — Italy: Cimolais. — Neth- erlands: Amsterdamse Bos; Bunde; Chaam; Dene- kamp; Hilversum; Hoogerheide; Oostvoorne; Ulven- hout; Winterswijk. — Yugoslavia: Bezdan. 11. Ectoedemia (Ectoedemia) turbidella (Zeller, 1848) (figs. 46, 100, 161, 162, 184, 242, 291, 362, 431, 432, 476, 519) Nepticula argyropeza var. turbidella Zeller, 1848: 321, 322. Syntypes, Poland: Glogow (Glogau), Zeller (depository unknown) [not examined]. [no genus] argyropeza; Herrich-Schäffer, [1853]: pl. 106 figs. 838, 839; [1854]: pl. 114 fig. 930 [misi- dentification]. Nepticula argyropezella Herrich-Schaffer, 1855: 357. (replacement name for turbidella Zeller). Nepticula populi-albae Hering, 1935: 7. Lectotype ? (here designated), Germany: Berlin, Tiergarten, 22.11.1933, M. Hering. Populus alba, N 4058, coll. Hypon., M. Hering, Genitalia slide on pin (MHUB) [examined]. Stigmella marionella Ford, 1950: 39, fig. Holotype 6, England: Stanmore, Middlesex, v, L. T. Ford (BMNH) [not examined]. [Nepticula argyropeza; Frey, 1857: 398—400, partim, misidentification |. Nepticula turbidella; Wocke, 1871: 339; 1874: 103; Heinemann & Wocke, 1877: 766; Sorhagen, 1886: 310; Rebel, 1901: 227; Meess, 1910: 480; Peter- sen, 1930: 76, fig. 115 (d genitalia); Hering, 1935: 7:Szöcs, 1965: 85. Stigmella turbidella; Klimesch, 1951: 64; Gerasimov, 1952: 265, 266; Klimesch, 1961: 762; Lhomme, 1963: 1202. Stigmella (Dechtiria) turbidella; Hering, 1957: 811, fig. 488a (mine). Dechtiria turbidella; Vari, 1950: 182, 184, figs. 9, 10 (8, 2 genitalia); Emmet, 1970a: 37—41, figs. (d genitalia, mine); 1971: 242, 243. Trifurcula (Ectoedemia) turbidella; Johansson, 1971: 245. Ectoedemia turbidella; Bradley et al., 1972: 3; 32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Borkowski, 1972: fig. 6 (d genitalia); Emmet, 1976: 189, pl. 12 fig. 36, pl. 7 fig. 2; van Nieuker- ken, 1982: fig. 2 (4 genitalia). Trifurcula turbidella; Karsholt & Nielsen, 1976: 18. Stigmella populialbae; Gerasimov, 1952: 252. Ectoedemia populialbae; Borkowski, 1975: 495. Diagnosis: see diagnosis of hannoverella for the differences between it and turbidella. The male genitalia resemble those of klimeschi, but can be recognised by the shape of the valva, with tooth-shaped tip in turbidella, and the asymmetric aedeagus in klimeschi. The female genitalia are very characteristic with the pointed ovipositor, and the long and broad apophyses. Description. Male. Forewing length 2.8—3.68 mm (3.34 + 0.21, 30), wingspan 6.2—8.4 mm. Head: frontal tuft light yellowish-orange or yellowish och- reous to dark fuscous; collar slightly lighter. Antenna with 46—59 segments (54.5 + 3.1, 23). Thorax blackish fuscous, with scattered white scales and sometimes a white tip; forewings blackish fuscous with a variable pattern of yel- lowish white spots: usually a medial costal and opposite dorsal spot; basal half with many scat- tered white scales, often forming a small discal spot halfway between wingbase and costal spot, and a basal spot along dorsal margin, usually giving a lighter appearance than hannoverella. Hindwing with a yellowish hair-pencil of about one-fifth of hindwing length. Female (fig. 46). Forewing length 2.76—3.48 mm (3.12 + 0.20, 29), wingspan 6.0—7.8 mm. Head: frontal tuft yellowish orange, never fus- cous. Antennae with 27—32 segments (29.3 + 1.3, 21). Ovipositor very conspicuous, pointed. Male genitalia (figs. 100, 242, 291, 362). Cap- sule length 270—347 um (304.3 + 23.0, 13). Te- gumen produced into a widely rounded pseu- duncus. Gnathos (fig. 291) with central element short triangular, smooth. Valva (fig. 242) length 193—227 um (210 + 9.5, 9), widest at base, gradually narrowing; tip inwards curved, tooth- shaped, clearly demarcated from valva. Aedea- gus (fig. 362) 369-399 um (377.6 + 13.9, 10), very long and stout, with two pairs of promi- nent carınae: ventral pair at extreme posterior tip, basally connected, pointed, single or with two or more tips; dorsolateral pair more ante- riorly placed, longer than ventral carinae, strongly curved, dorsally connected, often with additional spines at base, often asymmetrical. Female genitalia (figs. 161, 162, 184, 431, 432). T7 without row of setae. T8 relatively nar- row, tapering posteriorly, with two groups of 6—15 setae (to 20 in Iranian specimens), with- out or with very few scales. Anal papillae nar- row, with 7—12 setae. Anterior apophyses widened in middle, especially in lateral view. Posterior apophyses widening towards anterior end. Vestibulum with vaginal sclerite, a dorsal spiculate pouch with many (about 100) single, equally spaced, spines; and a patch of densely packed pectinations near entrance of ductus bursae. Corpus bursae relatively small, 420— 660 um, covered with small pectinations, except in anterior part, partly in concentric bands around signa; signa slightly dissimilar in length (not in shape), longest 219—283 um (263.6 + 23.7, 6), shortest 184—266 um (227.9 + 29.6, 6), 3.3—4.4 X as long as wide (data for speci- mens from Iran resp.: long signum 240—334 um; short 227—279, 2.7—3.4 x as long as wide). Ductus spermathecae with 2'4—3 con- volutions. Larva. Pale yellow. Sternite on prothorax on- ly. Ventral plates absent. This is the only Ectoe- demia s.str. species with dorsal as well as ven- tral calli. Biology. Hostplants: Populus alba L., P. canescens (Ai- ton.) Sm., only on the smaller leaves of older shoots of large trees, never on saplings. Material from Potsdam (leg. Hinneberg) is labelled with “Pop.nigr.”, but this is probably incorrect. Mine (figs. 476). Egg deposited on side of petiole, about 1/2—2 cm from leaf base. Mine first straight gallery in petiole, causing swelling; final instar larva makes triangular blotch be- tween first lateral vein and leaf margin, or less often between midrib and first lateral vein; frass deposited in two lateral lines, leaving passage for larva, which can withdraw itself in petiole. Live history. Univoltine. Larvae start feeding probably in summer, mature larvae can be found in October and November, usually later than hannoverella, often in green islands in fall- en leaves. The larva usually feeds in the night. Adults in May-June, or April in the South. Distribution (fig. 519). Widespread. In Scandinavia in southern Swe- den and Denmark only, very local in the ex- treme east of England, locally abundant throughout central Europe. Some scattered re- cords are known from southern Europe: Spain, Sicily. Also in North Iran (see remarks). Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 33 Remarks. Zeller (1848) described turbidella as a variety of argyropeza, as follows: “?Var. c. major; stri- gula ex costa prope basim obliqua dorsoque basali albidis, ceterum ut. b. Turbidella Z. in lit.”. Further (p. 322) he said that he believed it to possibly be a separate species. Unfortunately there is no specimen in the Zeller collection ın BMNH that can be identified as a syntype of turbidella. His description can, however, only refer to turbidella or hannoverella. Borkowski (1975) referred to turbidella types, examined by Johansson. However, Johansson (verbal comm.) only saw one specimen (genitalia slide BMNH 20537) which actually is argyropeza, and was sent by Mann in Vienna to Zeller in 1856, too late to be a turbidella syntype. Hence, the syno- nymy of turbidella with argyropeza by . Borskowski (l.c.) is unjustified. Ever since 1848, turbidella Zeller has been used for the species mining on Populus alba, with white scales in the basal half of the wing. This is not contradicted by the description, and thus its identity is firmly established. It is therefore not necessary to se- lect a neotype here. Unfortunately Herrich-Schaffer interchanged the names turbidella and argyropeza, although he knew exactly what Zeller meant by both names. Thus turbidella Herrich-Schaffer is a different species from turbidella Zeller, which was named by him’ first argyropeza Herrich- Schaffer (1853) and later argyropezella. Nepticula populi-albae Hering was described on the basis of a different head colour only, but since this is a very variable character, even with- in one population, it does not justify a separate identity. The fine series of turbidella collected by F. Kasy in Iran, consisting of only females, shows some slight differences from the European form in the genitalia. The most remarkable are that segments 8 and 9 are wider (fig. 432) and there is an oblique, hyaline bar in the vestibulum of all specimens examined (fig. 184). See also the measurements above. A sound taxonomic con- clusion about these specimens cannot be made without examination of males from the same re- gion, and preferably a study of the biology. The Iranian population is certainly not parthenoge- netic — as in argyropeza — for spermatophores were found in several of the bursae examined. Material examined: 175 8, 167 2. — Austria: 3 9, Hundsheimer Berg (near Hainburg), 15—16.v.1975, F. Kasy; 2 2, Klosterneuburg, Kuhau, 11.v.1915 and 14.111.1938, Preissecker; 1 d, Klosterneuburg, Krit- zendorfer Au, 16.111.1938, Preissecker; 1 2, Linz, 2.v.1910, Knitsche (NMW); 1 6,1 2, Linz, el. il iv.1936, Klimesch (NMW, ZMA); 1 2, Traun, e.l. 22—30.111.1936, J. Klimesch (ZMA); 1 2, Wien, Prat- er, 24.iv.1904; 13 6, 14 2, Wien, Prater, el. 4.iv— 21.vi.1984, E. J. van Nieukerken (ZMA); 1 d,1 9, Wien, Aspern, e.l. 27.v.1934, Koschabek; 1 ©, Lobau (Wien), 10.v.1908, Zerny (NMW). — Belgium: 3 9, Berg, 19.v.1945, L. Legiest; 2 d, Jette (Brussel), 28.iv.1945, L. Legiest; 1 dé, Laeken (Brussel), 13.v.1944, L. Legiest (IRSN). — Denmark: 1 6, 1 ?, Stigsnaes, 23.vi.1955, N. L. Wolff (MHUB). — Ger- many, West: 3 6,1 9, Bavaria, A. Schmid (RMNH); 1 6, 3 2, Braunschweig, Heinemann (MHUB). — Germany, East: 2 d, 1 © (lecto- and paralectotypes populialbae), Berlin, Tiergarten, el. 19—22.11.1933, Hering; 14 d, 6 ©, Berlin, Tiergarten, el. 11.1934, Hering; 16 d, 27 ©, Berlin, Botanische Garten, 11— 12.15.1948 and 17—28.11.1952, Hering (MHUB); 7 d, 9 ©, Potsdam, 28.1—10.11.1895, Pop. nigra (sıc!), Hinneberg (MHUB, ZMA, ZSM). — Great Britain: 2 3,2 2, Loughton: Epping Forest, el. 10—12.v.1980, Bryan & van Nieukerken (ZMA). — Iran: 10 ©, Ke- redj N., 27.iv.1970, Exp. Mus. Vind. (NMW). — Netherlands: 2 6, 1 9, Leiden, Leidse Hout, el. 22— 23.iv.1981, E. J. van Nieukerken; 9 d, 4 9, Oost- voorne, Mildenburg, e.l. 10—20.v.1983, Boomsma & Alders (ZMA); 1 6, Oostvoorne, 3.v.1981, Huisman (coll. Huisman); 1 4, Oostmaerland, 12.v.1974, G. Langohr, 28, Overveen, 10.v.1927 and 3.vi.1942, Bentinck; 37 d, 179, Santpoort, 1944—1948, Vari, Helmers, Doets (RMNH, ZMA); 1 6, 2 9, Santpoort N., Duin- en Kruidberg, e.l. 19.v.1983, Boomsma & Alders; 2 3, 2 2, Schinveld, 16.v.1976, G. Langohr; 39 8,23 2, Wijlre, 19.v.1974 and 22.v.1977, G. Lang- ohr (ZMA). — Poland: 5 6, 3 ©, Wroclaw (Breslau), el. i.1864 [Wocke] (MHUB, RMNH, ZMA); 1 ?, Wroclaw (Breslau), 19.v.1912 (NMW); 1 à, Silesia, Wocke (MHUB). — Spain: 2 6, Granada, 21—22.1v. 1883, Staudinger (MHUB); 1 d, 19 ©, Teruel, Valde- tormo, 8.v.1978, C. Gielis (coll. Gielis). — Yugosla- via: 6 ©, 2 km w. of Bezdan (Vojvodina), valley of Danube, el. 27.iv.—7.v.1984, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. — Austria: Muhlleiten (Grossenzersdorf); Wien, Prater. — France: Schirmeck. — Great Britain: Loughton, Epping Forest. — Netherlands: Santpoort. — Yugoslavia: Bezdan. 12. Ectoedemia (Ectoedemia) klimeschi (Skala, 1933) (figs. 47, 101, 163, 164, 243, 292, 363, 400, 401, 433, 478, 541) Nepticula klimeschi Skala, 1933: 31. Syntypes, Aus- tria: Linz, Donauauen, Populus alba, mines 1931, e.l. 1932, J. Klimesch (ZSMK, MHUB) [exam- ined]. Stigmella (Fomoria) niculescui Nemes, 1970: 33—35, figs. 1, 2. Holotype d, Rumania: Itcani (Suceava), 34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 16.iv.1966, 1. Nemes, Genitalia slide 1182 (coll. Nemes) [not examined] Syn. nov. [Nepticula argyropeza; Petersen, 1930: 78, fig. 122 (d genitalia) misidentification]. Stigmella klimeschi; Gerasimov, 1952: 244, 245; Klı- mesch, 1961: 763. Nepticula klimeschi; Hering, 1935: 7; Szöcs, 1965: 85. Stigmella (Dechtiria) klimeschi; Hering, 1957: 811 (mine). Ectoedemia klimeschi; Borkowski, 1975: 495. Diagnosis: females are externally almost inse- parable from E. argyropeza, only the larger number of antennal segments (34—38 in klimes- chi, 26—32 in argyropeza) being diagnostic. Fe- male genitalia can be separated from argyropeza by the signa which are longer in klimeschi, at least always longer than the shortest signum of argyropeza. There is some resemblance to the species of E. albifasciella-complex, but the latter have the costal spot always nearer the wing base and lack the hair-pencil in the male. See key for differences with suberis. Description. Male (fig. 47). Forewing length 2.76-3.6 mm (GOE) 0 272) win espan 16.032 main: Head: frontal tuft and collar yellowish orange. Antennae with 49—58 segments (52.8 + 3.2, 13). Thorax and forewings blackish fuscous, slightly irrorate by lighter scale-bases; a medial dorsal and costal white spot, opposite, usually widely separate; dorsal spot sometimes extend- ing along dorsal margin towards base. Hind- wing with yellowish hair-pencil of Y%—¥, hindwing length. Female. Forewing length 3.0—3.08 mm (3.05 + 0.03, 5), wingspan 6.7—6.8 mm. Antennae with 34— 38 segments (35.1 + 1.3, 15). Male genitalia (figs. 101, 243, 292, 363, 400, 401). Capsule length 292—321 um (307.7 + 11.9, 5). Tegumen produced into a widely rounded pseuduncus. Gnathos (fig. 292) with relatively long, triangular central element. Valva (fig. 243) length 214-236 um (226.3 + 8.2, 5), widest at base, gradually narrowing into trian- gular tip, not demarcated from valva. Aedeagus (figs. 363, 400, 401) 390—411 um (405.4 + 8.9, 5), very long and stout, markedly asymmetrical, posteriorly curved at right-hand side; with two pairs of prominent carinae: ventral pair at ex- treme posterior tip, basally connected, pointed, single; dorsolateral pair more anteriorly placed, longer than ventral carinae, strongly curved, dorsally connected, often with additional spine at base, which is larger in left process, asymme- trical. Female genitalia (figs. 163, 164, 433). T7 without row of setae. T8 wide, trapezoid, with two lateral groups of scales and many setae (13—20 at least). Anal papillae with 9-11 setae. Vestibulum with vaginal sclerite, a dorsal spicu- late pouch with many (more than 60) single, equally spaced, denticles; and a patch of densely packed pectinations near entrance of ductus spermathecae. Corpus bursae 660—715 um, covered with small pectinations, partly in con- centric bands around signa; signa almost simi- lar, 373—416 um (394.3 + 12.5, 8), 3.544 x as long as wide. Ductus spermathecae with 2%—3 convolutions. Larva. Pale yellow. Prothorax and segment 10 with sternites. Ventral plates absent. Biology. Hostplant: Populus alba L., on saplings and large lobed leaves of young branches on trees. When sympatric with turbidella, always on dif- ferent leaves, but sometimes on the same branch. Mine. (fig. 478). Egg on petiole, but almost impossible to find, between long hairs. Mine first straight gallery in petiole, causing it to swell. In final instar larva enters leaf, and makes blotch, usually not between veins, but incorpo- rating vein or midrib in middle of mine; frass in two lateral lines, leaving passage for larva, which can withdraw itself in petiole. Sometimes the larva feeds so long in the petiole, that there is hardly a mine in the lamina. E. klimeschi does not cause such conspicuous green islands as the related species. Life history. Univoltine. Larvae probably start feeding in summer, mature larvae can be found in October and November. Adults in June and July. Distribution (fig. 541). East and Southeast Europe, especially com- mon in Danube bassin, from West Germany to Rumania. Also recorded from East Germany, Poland, Switzerland and northern Italy. Remarks. The types from Skala’s collection are lost, but syntypes are still extant in other collections. I have examined syntypes from Berlin, but it would be more appropriate to select a lectotype from Klimesch’s collection. Previously this spe- neuburg, Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 35 cies has been confused with argyropeza, and probably most records of argyropeza males re- fer in fact to klimeschi. Stigmella niculescui Nemes is undoubtedly a synonym of klimeschi, the genitalia figure shows the characteristic valvae. The figure, however, is completely symmetrical, whilst the genitalia are asymmetrical. This species was previously believed to occur only in the leaves of saplings, but in 1983 we were able to find it also on similarly shaped leaves on large trees. Sometimes they were even found on the same branches as turbidella, but always in the lobed leaves. Material examined: 20 4, 19 2. — Austria: 1 9, Gramatneusiedl, Furbachwiesen, 11.vi.1982, F. Kasy; 2 d, 1 2, Hundsheimer Berg (near Hainburg), 13.vi.1979 and 8.vı1.1980, F. Kasy; 1 d, 1 ©, Kloster- Kritzendorfer Au, el. 15—16.1v.1938, Preissecker; 1 d, Klosterneuburg, Kuhau, el. 19.1v.1937, Preissecker; 1 ©, Klosterneuburg, Ziege- lofen, e.l. 17.v.1937, Preissecker (NMW); 1 2, Klos- terneuburg, Rollfähren, el. 19—21.v.1984, J. J. Boomsma & E. J. van Nieukerken (ZMA); 1 6, 1 2 (syntypes), Linz, Donauauen, el. 6.1v.1932, J. Kli- mesch (MHUB); 4 6, 2 ©, Linz, Donauauen, e.l. 21.1v.—2.v.1934, J. Klimesch (MHUB, NMW); 1 6, Linz, 12.v.1974, J. Klimesch; 4 6, 1 ©, Linz, Holz- heim, el. 3.vi.1980, 30.1v—12.v.1981, J. Klimesch; 2 6, 3 2, Wien, Prater, el. 15.v—19.vi.1984, E. J. van Nieukerken (ZMA). — Germany, Fast: 2 d, Bautzen, | e.l. 20—22.iv.1949, J.:Klimesch (ZMA). — Hungary: 1 4, Magyaresisatre, el. 12.1v.1917; 1 6, Nagy Nyir near Kecskemet, 4.v1.1914 (NMW). — Yugoslavia: 7 2,2 km W. of Bezdan (Vojvodina), valley of Danube, el. 5.„—12.v1.1984, J. J. Boomsma & E. J. van Nieu- kerken (ZMA). Mines. — Austria: Klosterneuburg; Linz (leg. Kli- mesch); Wien, Prater. — Germany, West: Munchen, Isarauen, 2.x1.1949, Groschke (BMNH). — Yugosla- via: Bezdan. Additional record: Italy: Piemonte, Poggio d’Aras- co, 9.vi.1977, Baldizzone (figure of d genitalia by Klimesch examined). 13. Ectoedemia (Ectoedemia) argyropeza (Zeller, 1839) (figs. 48, 165, 166, 434, 521) Lyonetia argyropeza Zeller, 1839: 215. Lectotype 9 (here designated) Poland: Silesia, Glogéw (Gross Glogau), 183., Zeller, Walsingham coll. 1910— 427; 101291, Genitalia slide BM 22611 (BMNH) [examined]. Nepticula apicella Stainton, 1854: 300. Lectotype ® (here designated), England: Beckenham, palings, 20.v.[18]51, Stainton, S 327/57, Genitalia slide 22610 (BMNH) [examined]. (Synonymised by Heinemann & Wocke, 1877). [No genus] turbidella Herrich-Schäffer, [1853]: pl. 106 fig. 837 [nomenclatorially unavailable]. Nepticula turbidella Herrich-Schaffer, 1855: 357, nec Zeller. Syntypes, Austria: Wien (depository un- known) [not examined]. Nepticula argyropezella Doubleday, 1859: 36 (unjus- tified emendation). Nepticula turbulentella Wocke, 1861: 129 (replace- ment name for N. turbidella Herrich-Schaffer nec Zeller). Nepticula simplicella Heinemann, 1862: 319, 320. Lectotype 2 (here designated), Germany: [Wolf- enbüttel], Buchheister (specimen painted... by R. Johansson) (Niedersächsisches Landesmuseum, Hannover) [examined by R. Johansson]. Syn. nov. Nepticula argyropeza ab. morosella Steudel & Hof- mann, 1882: 244. Nepticula argyropeza ab. houzeaui Dufrane, 1942: 11. Lyonetia argyropeza; Tengström, 1848: 152. Nepticula argyropeza; Zeller, 1848: 320, 321; Stain- ton, 1851: 11; 1854: 300; Frey, 1857: 398—400 [partım]; Stainton, 1859: 433 [partım, larva only]; 1862: 188—195, pl. 9 fig. 2 [partim, larva only]; Heinemann, 1871: 221; Nolcken, 1871: 795—797; Wocke, 1871: 339; 1874: 103; Heinemann & Wocke, 1877: 768; Sorhagen, 1886: 311; Tutt, 1899: 327—330; Rebel, 1901: 228; Meess, 1910: 481; Sorhagen, 1922: 57, pl. 4 fig. 66; Meyrick, 1928: 863; Hering, 1935: 7; Klimesch, 1936: 210; Szócs, 1965: 84. Nepticula apicella; Frey, 1857: 400, 401; Stainton, 1859: 433; Wocke, 1871: 339; Meyrick, 1895: 726. [Nepticula turbidella; Frey, 1857: 401, 402. Misiden- ufication]. Nepticula simplicella; Wocke, 1871: 340; Heinemann & Wocke, 1877: 770; Rebel, 1901: 228; Meess, 1910: 481. Stigmella argyropeza; Klimesch, 1951: 64; Gerasi- mov, 1952: 227; Klimesch, 1961: 763; Lhomme, 1963: 1205; Borkowski, 1969: 107. Stigmella (Dechtiria) argyropeza; Hering, 1957: 811 (mine). Dechtiria argyropeza; Emmet, 1971: 243, 244. Trifurcula (Dechtiria) argyropeza; Johansson, 1971: 245. Ectoedemia argyropeza; Bradley et al., 1972: 3; Borkowski, 1975: 494; Emmet, 1976: 189, pl. 7 fig. 4, pl. 12 fig. 35. Ectoedemia (Ectoedemia) 1972: fig. 11 (venation). Trifurcula argyropeza; Karsholt & Nielsen, 1976: 18. argyropeza; Borkowski Diagnosis: only females are known, which can easily be confused with klimeschi, see diag- nosis for that species. Description. Female (fig. 48). Forewing length (2.08) 2.6— 3.16 mm (3.16 + 0.25, 39), wingspan (4.5) 5.0— 36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 6.8 mm. Head: frontal tuft and collar yellowish orange. Antennae with 26—32 segments (29.0 + 1.7, 23). Thorax and forewings blackish fus- cous, slightly irrorate by lighter scale basis; a medial dorsal and costal white spot, opposite, usually widely separate; dorsal spot sometimes extending along dorsal margin towards base. Female genitalia (figs. 165, 166, 434). T7 without row of setae. T8 wide, trapezoid, with two lateral groups of scales and many setae (8— 12 at least). Anal papillae with 5-9 setae. Vesti- bulum with vaginal sclerite, a dorsal spiculate pouch with many (about 70) single, equally spaced denticles; and a patch of densely packed pectinations near entrance of ductus spermathe- cae. Corpus bursae 495—660 um, covered with small pectinations, partly in concentric bands around signa; signa slightly dissimilar, longest 270—394 um (325.4 + 42.0, 14), shortest 240— 351 um (307.3 + 35.4, 14), 3.44.3 X as long as wide. Ductus spermathecae with 2/2—3 convo- lutions. Larva. Pale yellow. Prothorax and segment 10 with sternites. Ventral plates absent. Biology. Hostplant: Populus tremula L. Mine. Egg on side of petiole, about 1 cm from leaf base. Mine first straight gallery in petiole, causing it to swell, later blotch in lamina be- tween midrib and first lateral vein; frass in two lateral lines, leaving passage for larva, which of- ten hides in petiole; mine similar to turbidella. Life history. Univoltine. Larva starts feeding early, from July, mature larvae can be found from early September to November, often in green islands in fallen leaves. The larva feeds usually in dark only. Time of completing larval cycle largely depends on age of leaf: when leaf falls in September the larva will be full-grown a long time before larvae in leaves still on the tree. This probably applies as well to the related spe- cies. Adults in May and June. Distribution (fig. 521). Widely distributed in Europe, and often very abundant. Not yet recorded from Ireland, Iberi- an Peninsula and south of Po valley, North Yu- goslavia and Rumania. Remarks. Although a distinct species, E. argyropeza has been the subject of much nomenclatorial confusion. I have designated as lectotype the specimen in the Zeller collection, which had been labelled holotype by Durrant. Herrich- Schaffer (1853, 1855) was aware of the differ- ence between argyropeza and turbidella, but in- _terchanged these names deliberately and thus renamed argyropeza as N. turbidella. This in- correct use has, however, only been followed by Frey (1856, 1857). Stainton correctly described the biology of argyropeza, but mistook the adult of albifasciella for argyropeza (see under albifasciella). He therefore had to rename the real argyropeza, and gave it the name apicella. R. Johansson examined the types of N. sim- plicella Heinemann and found they were just uniformly coloured examples of argyropeza. By courtesy of Mr. Johansson I designate here the lectotype that he selected but did not publish. E. argyropeza is a parthenogenetic species, of which males are unknown. Reported males be- long to either albifasciella or klimeschi. We have several times bred larvae from single females, — which therefore corroborates their absolute par- thenogenetic reproduction. Wilkinson & Scoble (1979) reported the species also from Canada and the USA, where it is parthenogenetic as well. Study of Canadian material showed that there is not much difference in morphology or allozyme pattern (Menken, in preparation) be- tween them and the European populations. It is therefore likely that the North American popu- lations are the offspring of recent introductions which may not warrant subspecific status. Material examined: 169 9. — Austria: 1 9, Gumpoldskirchen, Glaslauterriegel, 17.v.1983, F. Ka- sy (NMW); 1 2, Linz, el. 17.11.1932, J. Klimesch (MHUB); 12, Waldburg (near Freistadt)), el. 13.1.1921, Knitschke; 3 2, Wien, Haschberg, e.l. 12—18.11.1937, Preissecker; 1 2, Wien, Prater, 1867 (NMW); 22, no further data (RMNH). — France: 1 2, Malesherbes (Loiret), 8.v.1955, Buvat; 1 9, Puy Saint Vincent (Hautes Alpes), 6.vi.1965, Buvat (coll. Buvat). — Germany, West: 3 9, Braunschweig, Heinemann (MHUB); 1 ©, Freiburg (MHUB); 1 9, Heidelberg, Ziegelhausen, 17.v.1976, W. Speidel (coll. Speidel). — Germany, East: 30 ®, Berlin, e.l. v. Hering; 8 ©, Nauen, el. 24.11—2.11.1924, Hering (MHUB); 9 ©, Potsdam, el. 13—19.11.1893, Hınne- berg (MHUB, ZMA). — Great Britain: 6 ©, Berley, Kent, 15.v.1947, S.N.A. Jacobs (ZMA); 3 ©, (lecto- and paralectotypes of apicella), Beckenham, palings, 20—25.v.1851, Stainton (BMNH). — Italy: 2 ©, Na- turno (Bolzano), 2 km SE, N. slope, 800 m, e.l. 4.v.1984, J. J. Boomsma (ZMA). — Netherlands: 83 2 from following localities: Berg en Dal; Denekamp; ’s-Graveland; Groote Peel; Hilversum; Overveen; Winterswijk; Zwanewater (RMNH, ZMA, coll. Kos- ter). — Poland: 6 2, Wroclaw (Breslau), e.l. 15— VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 37 16.v.1858, 1863, [Wocke] (MHUB); 1 2 (Lectotype, see above). — No data: 1 9, e.l. 24.11.1866 (MHUB). Mines. — Austria: Peggau. — France: Barr. — Germany, West: Birresborn; Blankenheim; Wies- baum. — Germany, East: Berlin, leg. Hering (BMNH). —Great Britain: Earls Colne (Essex). — Hungary: Budapest. — Italy: Naturno. — Nether- lands: many localities. — Yugoslavia: Fuzine, SW of Delnice. The Ectoedemia preisseckeri group 14. Ectoedemia (Ectoedemia) preisseckeri (Klimesch, 1941) (figs. 49, 102, 167, 168, 244, 293, 364, 402, 435, 479, 540) Nepticula preisseckeri Klimesch, 1941: 162—168, figs. 1—10, pl. 16. Lectotype d (here designated) Aus- tria: Klosterneuburg, Kritzendf. Au, e.l. 2.v.1939, Preissecker, Ulm., Genitalia slide MV 12214 (NMW) [examined]. Stigmella preisseckeri; Hering, 1957: 1092, fig. 698, 705b (mine); Klimesch, 1961: 760. Ectoedemia (Dechtiria) preisseckeri; Klimesch, 1975c: 11,3 figs. (d genitalia, mine). Ectoedemia preisseckeri; Borkowski, 1975: 493. Diagnosis: externally almost inseparable from albifasciella-complex, see key-characters. Male genitalia characterised by two pairs of similar, curved carinae and triangular gnathos. Female genitalia differ by combination of pectinate bur- sa and slightly dissimilar signa, which are much shorter than in albifasciella-complex. Description. Male (fig. 49). Forewing length 5.6—6.0 mm (2.63 + 0.09, 6), wingspan 5.6—6.0 mm. Head: frontal tuft and collar yellowish orange to ferru- ginous. Antennae with 36—39 segments (37.2 + 1.2, 6). Thorax and forewings blackish fuscous, thorax without white scales at tip of mesoscu- tum and tegulae; forewing with yellowish white, not shining spots: one dorsal in middle, and one costal before middle, sometimes united to form fascia. Hindwing without hair-pencil but with costal bristles. Female. Forewing length 2.56—2.68 mm (2.62 + 0.06, 5), wingspan 5.7—6.0 mm. Anten- nae with 27—31 segments (29 + 1.6, 5). Male genitalia (figs. 102, 244, 293, 364, 402). Capsule length 257—317 um (4). Tegumen pro- duced into broadly triangular pseuduncus. Gnathos (fig. 293) Sith central element triangu- lar, pointed. Valva (fig. 244) length 214 um (3), widest at base, inner margin serrate by promi- nent setal sockets, tip rounded; posterior mar- gin with a notch, in ventral view suggesting a double tip. Aedeagus (fig. 364, 402) 330—334 um (4), with a dorsal and dorsolateral pair of strong, curved carinae of same length, dorsal pair often overlapping; aedeagus slightly con- stricted. Female genitalia (figs. 167, 168, 435). Ab- dominal tip narrow. T7 with a row of 6—12 se- tae along posterior margin. T8 approximately quadrate, with two groups of 1-4 setae, without scales. Anal papillae with 12—19 setae. Vestibu- lum with vaginal sclerite, a dorsal spiculate pouch with many spines, both single and in rows, and a dense patch of pectinations near en- trance of ductus spermathecae. Ductus bursae densely covered with pectinations. Corpus bur- sae 550—790 um, covered with small pectina- tions, except anterior part; signa ovoid, slightly dissimilar in length, longest 369—441 um (3), shortest 330—394 um (3), 2.43.0 X as long as wide. Ductus spermathecae with 21/5—312 con- volutions. Larva. Whitish, with distinct ganglia. Penulti- mate stages with 12 dark brown ventral plates, which are shed during final instar. See detailed description by Klimesch (1941). Biology. Hostplant: Ulmus spp. Mine (fig. 479). Egg on either side of leaf, on a vein. Early mine narrow, much contorted gal- lery, with frass in widely separated pellets, then abruptly widening into elongate blotch, with blackish frass concentrated in basal half or at margins, often absorbing early gallery. Life history. Univoltine. Larvae in Septem- ber—Oktober. Adults probably in May-June (reared in April—June). Distribution (fig. 540). Only known from the Danube valley, near Vienna and Budapest, although not always near the river. Material examined: 84, 5 2. — Austria: 2 d, Bad Deutsch Altenburg, Pfaffenberg, 3 km SW Hainburg, e.l. 21.vi.1984, E. J. van Nieukerken (ZMA); 1 4,1 2 (paralectotypes), Klosterneuburg, e.l. 23.iv.1939, J. Klimesch; 2 d, 1 © (lecto- and paralectotypes), Klos- terneuburg, Kritzendf. Au, e.l. 1—3.v.1939, Ulm., Preissecker; 1 9, Klosterneuburg, Kuhau, el. 7.v.1939, Preissecker; 1 6, Wien, L.-Enzersdorf, e.l 19.v.1918, Ulm, Preissecker (NMW). — Hungary: 2 3, 2 9, Budapest, Kamaraerdö, el. 19—20.v.1975, Ulmus camp., J. Szöcs (TMAB). Mines. — Austria: Bad Deutsch Altenburg (Hain- burg); Wien, Prater. 38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 The Ectoedemia suberis group The species of this group feed on Quercus species, and make blotch mines. Except aegilo- pidella, they form a relatively uniform group of fasciate moths, with conspicuous hair-pencil in male, and often a hairy abdominal tip in female. Male genitalia have large curved valvae, one pair of single carinae, and a simple gnathos. Female genitalia are characterised by weak development of vaginal sclerite and spiculate pouch, a globular bursa, covered with pectina- tions and wide, similar, oval signa. The larvae are invariably green and have no ventral plates. The European species all occur in the south- ern part and the group is probably also present in the Eastern Palaearctic area (E. chasanella Puplesis, 1984a). 15. Ectoedemia (Ectoedemia) caradjai (Groschke, 1944) (igs 5011035 178,245,2406,294,365,407,.436, 483, 526) Nepticula caradjai Hering, 1932: 16. [nomen nudum, description of mine only]; Toll, 1934b: 72 (record of mine). Nepticula caradjai Groschke, 1944: 118, figs. 3, 4. ? Holotype ©, [Italy: Sicilia, Taormina], 518 [e.l. 9.1x.1942, Quercus pubescens, F. Groschke] (SMNS) [examined]. Stigmella caradjai; Klimesch, 1951: 65, fig. 73; Gera- sımov, 1952: 232; Hering, 1957: 876, figs. 530, 538, 544 (mine); Klimesch, 1961: 762. Nepticula caradjai; Sz6cs, 1965: 87. Trifurcula (Ectoedemia) caradjai; Klimesch, 250, figs. 23, 24 (mine). Ectoedemia caradjai; Sz6cs, 1981: 211. ? Trifurcula (Ectoedemia) species; Klimesch, 1978, 250, 251, fig. 25 (mine). 1978: Diagnosis: male recognised by combination of fascia, basal white streak on forewing and white hair-pencil, female by same wing-pattern and hairy abdominal tip. Sometimes basal streak inconspicuous, then similar to larger suberis, but in male of caradjai hair-pencil not surrounded by special scales. See also leucothorax. Male genitalia characterised by shape of valva. Female genitalia separated from suberis by shape of sig- na. Description. Male. Forewing length 1.88—2.4 mm (2.19 + 0.15, 13), wingspan 4.2—5.3 mm. Head: frontal tuft yellowish to yellow mixed fuscous; collar yellowish. Antennae long, with 43—51 seg- ments (48.3 + 2.6, 11), scape with some brown scales. Thorax brown, with some white scales, especially at tip of mesoscutum and tegulae. Forewings fuscous, with a basal white streak along dorsum, sometimes joining fascia, sometimes inconspicuous, and a medial, almost straight fascia, sometimes broken. Hindwing with snowwhite hair-pencil of ‘4 hindwing length, not surrounded by special scales. Female (fig. 50). Forewing length 2.32—2.68 mm (4), wingspan 5.2—5.8 mm. Antennal seg- ments 30—32 (4). Male genitalia (figs. 103, 245, 246, 294, 365, 407). Capsule length 244—261 um (251.1 + 7.8, 5). Tegumen produced into small, but distinct, rounded pseuduncus (fig. 407). Gnathos (fig. 294) with narrow long central element, blunt at tip, with smooth margins. Valva (fig. 245) length 171193 m'(1187-7, #93, 5) mnermar- gin basally almost straight, gradually becoming strongly concave towards pointed tip; outer margin uniformly convex. Aedeagus (fig. 365) 274—291 um (282 + 7.0, 5), cärınae pointed, single, curved outwards. Female genitalia (figs. 173, 436). T7 with a crescent-shaped patch of at least 100, very long setae, appearing pectinate at large magnifica- tions (1000 x). In addition T7 + 8 covered with about 50 shorter, more widely spaced setae, T8 without scales. Anal papillae wide, each with about 40 setae. Vestibulum with vaginal sclerite, and an indistinct dorsal spiculate plate, with few spines. Corpus bursae 495—570 um, covered with pectinations, except in distal third; signa almost similar, 300— 394 um (6), 2.4—2.7 umes as long as wide. Ductus spermathecae with 31/,—4 inconspicuous convolutions. Larva. Green. Ventral plates absent. Biology. Hostplants: Quercus pubescens Willd. s.l., from which it has been reared most often. Mines recorded from: Q. frainetto Ten., Q. petraea L. s.l.. About occurrence on Q. infectoria Olivier and Q. coccifera L. see remarks. Mine (fig. 483). Egg on either surface, usually near or at margin. Early mine narrow contorted gallery up to 1.5 cm long, filled with frass, abruptly enlarging into roundish or elongated blotch with frass heaped near entrance, or in two lateral lines. Life history. Univoltine. Larvae from July to September, adults from late May to early July, earlier records refer to reared material. Klı- mesch (1978) supposed that a second generation | Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 39 occurred in Anatolia, since his July larvae gave rise to adults in August. Distribution. (fig. 526). In central and southern Europe, south and east of the Alps. Westernmost locality is in France. Not yet recorded, but to be expected from Czechoslovakia, Rumania and Bulgaria. Remarks. Most authors incorrectly attribute the name caradjai to Hering. Although he gave this name to the species, it is not available, since he de- scribed the mine only, after 1930 (Code, art. 13a, 16). Toll was the first who reared the adult, and some of his specimens are labelled as type, but he failed to describe the species. Thus, Groschke has to be regarded as the author, since he was the first who described caradjai. The collection of Groschke is in SMNS, but it is al- most useless, since his specimens bear only la- bels with a number. According to W. Speidel (pers. comm.) no diaries or notebooks belong- ing to Groschke could be traced to find the meaning of these numbers. However, when I borrowed all the Nepticulidae from this collec- tion, it was apparent that all specimens num- bered from 514 tot 573 belong to species, which were collected in Taormina, Sicily, during the war. We know this from Groschke (1944) and from the Hering Herbarium (BMNH), where many nepticulid mines collected by Groschke are to be found. It is furthermore notable, that the first (514) and last (573) number are repre- sented by species, which are typically mediter- ranean, i.e. Nepticula euphorbiella Stainton and N. groschkei Skala, therefore probably none of this series was collected elsewhere. There is one 2 specimen in this collection, labelled 518, which corresponds completely with Groschke’s description, and undoubtedly belongs to carad- jai, but unfortunately lacks the abdomen. Groschke only mentioned one specimen in his description, so with some reluctance, it 1s ac- cepted as the holotype of caradjai. It is not yet clear if caradjai is one variable species, or forms a complex comparable with subbimaculella. Klimesch reared some very sim- ilar specimens from the semi-evergreen Quercus infectoria and the evergreen Q. coccifera (Kli- mesch, 1978). These specimens differ slightly since they are smaller, but do not show diagnos- tic differences. Their measurements are therefore excluded from the above mentioned data, but follow here: 1. from GQ. infectoria. 3d: forewing length 1.84—2.04 mm (2), antennal segments 47— 48. Capsule 206 um, valva 171 um (fig. 246), aedeagus 244 um. 9 : forewing length 1.88—2.04 mm, antennal segments 31—32. Bursa 440 um, signa 227—270 um, 2.3—2.5 x as wide as long, less setae on T8 and anal papillae (about 20) (see fig. 174). 2. from Q. coccifera. 2: forewing length 2.2 mm, antennal segments 33. Bursa 570 um, signa 334—343 um, 2.8 X as long as wide, 30 setae on anal papillae; ductus spermathe- cae with 3 convolutions (fig. 175). More material is needed to check the constan- cy of these observations, and also especially to compare specimens reared from Q. pubescens on Rhodos. Material examined: 17 6, 7 2. — Austria: 1 d, Gumpoldskirchen, Glaslauterriegel, 4.v11.1976, Kasy; 2 8, Hackelsberg, N. of Neusiedlersee, 23.v1.1975, 29.vi.1977, Kasy; 1 d, Wien, Leopoldsberg, e.l. 26.v.1943, Q. pubescens, Preissecker (NMW). — Hungary: 1 6, Csopak, e.l. 24.v.1971, J. Szöcs; 1 2, Nagykovacsi, Remetehegy, el. 19.vi.1963, J. Szócs (TMAB). — Italy: 6 d, Monti Aurunci (Latina), 4 km NW Castelforte, 400 m, 22—23.vi + 1.vu.1969, R. Johansson (coll. Johansson); 1 d, Sitizano (Calabria), 450 m, 28.viii.1977, S. E. Whitebread (coll. White- bread); 1 2, ? Holotype (see above). — Turkey: 1 6, Anatolia, Kizilcahamam, 700 m, 31.vii—1.vi.1963, Arenberger (LNK); USSR: 2 d, 4 2, Babince, k. Rae (Reclelle), else 5s 388, 187 1eı— 1.iv.1939, Q. pubescens, S. Toll (IPAK, MHUB); 2 5, Scianka Hlody, p. Borszezów (Podolia), el. 25 26.11.1939, S. Toll (IPAK, MHUB). — Yugoslavia: 1 ?. Treschkaschlucht, near Skopje, 21—30.v1.1959, F. Kasy (NMW). Identity uncertain: 2 d, 4 2. — Greece: 2 d,3 9, Rhodos, Treas, e.l. 20—30.iv.1978, Quercus infecto- ria, J. Klimesch; 1 2, Rhodos, Trianta, e.l. 4.v.1974, Quercus cocafera, J. Klimesch (ZSMK). Mines. — On Quercus frainetto. — Greece: Oiti Oros (Fthiotis). On Quercus petraea s.l.. — Greece: W. Palaiokastron (Evritania). On Quercus pubescens. — Austria: Gumpoldskirchen; Hainburg: Hunds- heimer Berg. — France: Aix-en-Provence, leg. J. W. Schoorl; Viens (Vaucluse) (near Apt), leg. R. Buvat. — Greece: Evvoia, Dhirfis Oros; Oiti Oros (Fthio- tis); Voutonasi (Ioannina). — Italy: Abruzzi: Goia dei Marsi; Picinisco; Lazio: Veio; Sicilia, Taormina, leg Groschke (BMNH); USSR: Bendery (Tighina), leg. Hering (BMNH). Identity uncertain: on Quercus infectoria. — Greece: Rhodos, leg. Klimesch. 16. Ectoedemia (Ectoedemia) species (specimen 1843) (figs. 104, 247, 295, 366) 40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Material: 1 d: Spain: Aragon, Rubielos de Mora, 4.vii.1967, Arenberger, Genitalia slide VU 1843 (LNK). This specimen clearly belongs in the group near caradjai and suberis, but is almost certainly specifically different. Due to the bad condition of the specimen, however, I refrain from nam- ing it. It is most easily separated from the other species in the group by the ochreous brown hair-pencil, surrounded by brown lamellar scales. The genitalia are most similar to caradjat. Description. Male. Forewing length 2.44 mm, wingspan 5.4 mm. Antennal segments not countable. Worn specimen, wing pattern similar to suberis. Hindwing with ochreous brown hair-pencil, surrounded by brown lamellar special scales. Male genitalia (figs. 104, 247, 295, 366). Cap- sule length 257 um. Tegumen very broad, trun- cate. Gnathos with triangular central element (fig. 295). Valva (fig. 247) length 206 um, inner margin concave, outer margin strongly convex, tip pointed. Aedeagus (fig. 366) 304 um, carinae pointed, single. 17. Ectoedemia (Ectoedemia) suberis (Stainton, 1869) comb.n. (figs. 51, 105, 169, 170, 248, 296, 367, 408, 437, 480, 542) Nepticula suberis Stainton, 1869: 229. Lectotype d (here designated), France: Cannes, e.l., found dead, iii.[18]68, Q. suber, green larva, Stainton, Genitalia slide BM 22577 (BMNH) [examined]. Nepticula viridella Mendes, 1910: 165, pl. 7, figs. 6, 9. Syntypes, Portugal, prov. Beira Baixa, San Fiel, Mendes (depository unknown) [not examined] Syn. nov. Nepticula suberis; Wocke, 1871: 338; Rebel, 1901: 227; Meess, 1910: 479; Petersen, 1930; 71, fig. 101 (6 genitalia). Stigmella suberis; Gerasimov, 1952: 262; Hering, 1957: 868, fig. 539 (mine); Lhomme, 1963: 1196. Stigmella (Stigmella) suberis; Leraut, 1980: 48. Nepticula viridella; Hering, 1935: 373. Stigmella viridella; Gerasimov, 1952: 260; Hering, 1957: 867 (mine). Diagnosis: separated from caradjai by ab- sence of white basal streak on forewing, and presence in male of white lamellar scales, sur- rounding hair-pencil. The hair-pencil in male, and the dense group of long setae on the female postabdomen also separate suberis from haraldi and other similar oak-mining species. In male genitalia the shape of the valva is very character- istic. See also diagnosis for andalusiae. Description. Male. Forewing length 2.72—3.08 mm (2.95 + 0.09, 22), wingspan 6.5—6.8 mm. Head: frontal tuft yellowish orange to ferruginous; collar lighter. Antennae long with 49—60 short segments (54.9 + 3.3, 17). Thorax and forewing brown, irrorate with white; a medial almost straight dull white fascia. Hindwing with white hair-pencil surrounded by white special lamellar scales. Female (fig. 51). Forewing length 2.8—3.24 mm (3.05 + 0.10, 23), wingspan 6.4—7.2 mm. Antennal segments 37—43 (39.1 + 1.5, 18). Male genitalia (figs. 105, 248, 296, 367, 408). Capsule length.261—296 um (279.5 + 11.5, 9). Tegumen produced into broadly triangular, rounded pseuduncus (fig. 408). Gnathos (fig. 296) with long triangular central element. Valva (fig. 248) length 201—227 um (212.7 + 9.1, 8), basally broad with inner margin convex, below | middle suddenly narrowed and inner margin be- coming concave towards tip. Aedeagus (fig. 367) 343—394 um (375 + 18.0, 8), much longer than capsule, carinae single, pointed, slightly curved outwards. Female genitalia (figs. 169, 170, 437). T7 with a semicircular patch of 120—200 very long, smooth setae. T7 and 8 in addition with about 80—100 shorter setae, without scales. Anal pa- pillae with 29—37 setae. Vestibulum with vagi- nal sclerite and a spiculate pouch with hardly visible spines, without pectinations. Corpus bursae almost globular, 550—660 um; covered with minute pectinations; signa similar, 364— 437 um (417.4 + 40.0, 10), 2.3—2.4 x as long as wide. Ductus spermathecae with 4—4' distinct convolutions. Larva. Dirty green, with conspicuous brown ganglia. Ventral plates absent. Biology. Hostplants: Quercus suber L., Q. ilex L., Q. rotundifolia Lam, Q. coccifera L. and possi- bly Q. faginea Lam. Mine (fig. 480). Egg on leaf-upperside. Mine starts as contorted gallery filled with frass, later widening into large irregular blotch with the frass in basal half or in two lateral lines. Larva feeds only in upper parenchym layers. Life history. Univoltine. Larva feeds in win- ter, mainly from January to March, occasionally early April. Larva or pupa aestivates in cocoon, adult flies from July to early October, but some specimens from Marbella were taken in June. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 41 Distribution (fig. 542). Western mediterranean species, known from Iberian peninsula, France, Corsica, Sardınıa and North Africa. Not recorded from mainland i Italy. Remarks. In contrast with ilicis, no types of viridella . Mendes could be found in the De Joannis col- lection in Paris, but it does contain two speci- mens, labelled viridella, collected in Salamanca (Spain), probably by Mendes, who lived there after 1910 (Zerkowitz, 1946). These specimens are identical with suberis. Also Mendes’ descrip- tion does not give reason to believe that virıdel- la should be regarded as a distinct species, it is therefore synonymised here. Material examined: 33 gd, 33 ©. — France: 1 6, Lectotype (see above); 2 6d, Cannes, Ragonot (RMNH); 2 d, 4 9, Alp. Mar., Cannes, Constant (IRSN, MNHN, RMNH); 1 2, Collobriéres (Var), e.l. 3.1x.1981, Quercus suber, S. E. Whitebread (coll. Whitebread); 3 6, Corse, el. 29.vin + 6.1x.1906, Q. ilex, Chrétien (MNHN); 1 ©, Golfe Juan, Alp. marit., Constant (IRSN); 1 d, 3 9, « Nesp. » (? near St. ' Pons, dep. Hérault), 2.vii.1904, Chrétien; 3 d, St. Pons, 4.vii.1904, Chrétien (MNHN). — Italy: 3 6, Sardegna, Mt. Istiddi, 1.1x.1978, G. Derra; 1 d, Sar- degna, Bacu Trotu, Ortuabis, 800 m, 28.vi11.1978, G. Derra (coll. Derra); 1 4, Sardegna, prov. Nuoro, Vil- lanova-Strisaili 885 m, 7.vii.1983, J. Kuchlein (coll. Kuchlein). — Morocco: 1 d, Tanger, 2.v.1902, Wal- singham (BMNH). — Spain: 1 9, Albarracin (Arago- nia), el. 1x.1933, Quercus ilex, Hering (MHUB); 1 4, 1 2, Alcuescar, Caceres, 1.x.1983, C. Gielis (Coll. Gielis; EvN); 3 d, Andalucia, prov. Malaga, road to Ojen, 150 m, 12.vi.1981, E. Traugott-Olsen (ETO); 1 3,7 km N. Benahavis (Málaga), road to Ronda, 800 m, e.l. 21—22.viti.1984, Quercus coccifera, E. J. van Nieukerken (ZMA); 1 d, 1 2, La Vid (Burgos), 800 m, 23—28.1x.1965, H. G. Amsel; 1 &, Cataluna, Port Bou, 18—28.ix.1966, Arenberger (LNK); 1 ?, 4 km NE Igualeja, Serrania de Ronda (Malaga), 1100 m, e.l. 21—22.vi11.1984, Quercus rotundifolia, E. J. van Nieukerken; 1 d, 7 2, Marbella (Malaga), Casa y Campo, 100 m, el. 29.viii—29.x.1984, Quercus cocci- fera, E. J. van Nieukerken (ZMA); 1 d 2 2, Las Mur- tas (near Elche), Murcia, 23.ix.1983, C. Gielis (coll. Gielis, EvN); 1 d, 1 2, Salamanca, el. 25.viii., Q. ilex [Mendes], coll. de Joannis (as viridella) (MNHN); 1 2, San Roque, Cadiz, 29.ix.1983, C. Gielis; 4 d, 10 2, Sierra Blanca, 6 km N. Marbella (Málaga), El Mira- dor, 800 m, e.l. 13—28.viii.1984, Quercus suber + ro- tundifolia, E. J. van Nieukerken (ZMA). Mines. — On Quercus suber. — France: Collo- brieres, Var., leg. Whitebread; Plan d’Aups, Var., leg. Whitebread (coll. Whitebread). — Spain: prov. Mala- ga: Casares; Istan; Marbella; Serranıa de Ronda; Sier- ra Blanca, N. Marbella. — Tunisia: Jebel Abiod; Ain Draham. On Quercus ilex. — France: Corsica, Barbi- caja, leg. Buhr (BMNH). On Quercus rotundifolia. — Algeria: Aurés Mts, near Arris; Aures Mts, Dj. Chélia. — Spain: Sierra Almijara, N. Otivar; Sierra Blanca, N. Marbella; Serrania de Ronda. Identity un- certain: on Quercus faginea. — Spain: Istan. 18. Ectoedemia (Ectoedemia) andalusiae sp. n. (figs. 52, 106, 171, 172, 249, 297, 368, 409, 438, 481, 526) Type material: Holotype 2: Spain (Malaga): Marbella, Casa y Campo, 100 m, 8.11.1984, e.l. 17—18.v.1984, Quercus coccifera, VU no. 84043 KE, E. J. van Nieukerken, Genitalia Slide 1899 (ZMA). Paratypes, 4 6, 3 9. — Spain: 2 d, 1 2, Andalucia, prov. Malaga, Camino de (road to) Ojen, 150 m, 12.v1.1981, E. Traugott- Olsena ZN AERO) RS dem Aleviel Osos ©, Andalucia, prov. Málaga, Camino de (road to) Istan, 400 m, 4.v11.1973, E. Traugott-Olsen (ETO); 1 3, Marbella, Casa y Campo, ca 100 m, 18.1x.1982, E. Traugott-Olsen (ETO); 1 2, Pyr. Orient., Tolorin b. Martinet, 6.v11.1967, Arenberger (LNK). Mines examined: on Q. coccifera from type locality, mixed with E. suberis mines. Diagnosis: 2 separated from suberis by ab- sence of long setae on abdominal tip; from ha- raldı by straighter fascia and genitalia. d very similar to suberis, separated by ochreous-brown hair-pencil instead of white, and markedly shorter capsule with blunt and wide tegumen. Description. Male. Forewing length 2.44—2.72 mm (4): wingspan 5.4—6.2 mm. Head: frontal tuft and collar yellowish-orange. Antennae with 49-57 segments. Thorax and forewings brown, with medial, almost straight, constricted, dull-white fascia. Hindwing with ochreous-brown hair- pencil, surrounded by white lamellar scales as in suberis. Female (fig. 52). Forewing length 2.4—3.04 mm (4), wingspan 5.5—6.9 mm. Antennae with 35—38 segments. Male genitalia (figs. 106, 249, 297, 368, 409). Capsule length 223—261 um (4). Vinculum an- teriorly narrower than in suberis. Tegumen truncate, very broad, hardly produced into pseuduncus (fig. 409). Gnathos (fig. 297) with tiangular central element. Valva (fig. 249) length 193—210 um (4), basally broad with inner mar- gin convex, below middle suddenly narrowed 42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 and inner margin becoming concave towards tip. Aedeagus (fig. 368) 309—351 um (4), much longer than capsule, carinae single, pointed, slightly curved outwards. Female genitalia (figs. 171, 172, 438). T7 without long setae. T8 with two lateral patches of scales and 4—7 setae. Anal papillae wide, with 18—24 setae. Vestibulum with vaginal sclerite and a spiculate pouch with very few, small spines, without pectinations. Corpus bur- sae almost globular, 495—640 um; covered with minute pectinations; signa similar, 330—377 um (348.2 + 1.41, 8), 1.92.5 x as long as wide. Ductus spermathecae with 512 convolutions. Larva. As suberis. Biology. Hostplant: Quercus coccifera L., from which holotype was bred. Mine (fig. 481). Not differentiated from the mine of suberis. Life history. Adults taken in June, July and one male in September, larvae found ın January. Distribution (fig. 526). Only known from Spain. Remarks. This species is closely related to E. suberis, but the female shows several diagnostic fea- tures, especially in the abdominal tip. The holo- type was reared from a mixed sample of mines collected on Quercus coccifera, from which also suberis has been reared. The mines do not give any evidence of the presence of two species. 19. Ectoedemia (Ectoedemia) aegilopidella (Klimesch, 1978) comb. n. (figs. 53, 54, 107, 176, 250, 298, 369, 410, 439, 482, 546) Trifurcula (Ectoedemia) aegilopidella Klimesch, 1978: 269—271, figs. 65—69. Holotype d, Greece: Rhodos: Rodini, e.l. 17—30.iv.1973, Zucht nr. 1054, Quercus macrolepis, 22.1x.1972, J. Klimesch, Genitalia slide Kl. 4107 (ZSMK) [genitalia slide examined]. Diagnosis: very small species with a wing- span of less than 4.2 mm. Males with basal % of hindwing covered with brown special scales, as in heringella and terebinthivora, but separated from these two by presence of a hair-pencil in aegilopidella. Females very similar to terebin- thivora, but terebinthivora has a more yellow fascia. Male genitalia very characteristic and di- agnosed by small size, wide capsule, gnathos and tegumen. Female genitalia characterised by absence of group of many long setae and small and short signa. Description. Male (fig. 53). Forewing length 1.80—1.92 mm (2), wingspan 4.0—4.2 mm. Head: frontal tuft and collar yellowish white. Antennae with 35—37 segments (2). Thorax and forewings ochreous-brown, with a medial, often ill-de- fined, straight fascia, colour yellowish white. Underside of forewing with a group of brown androconial scales in distal half, and a group of short, yellowish-white lamellar scales near cos- tal retinaculum. Hindwing with a yellowish- white hair-pencil of ‘4 hindwing length; basal % covered with brown lamellar, special scales. Female (fig. 54). Forewing length 1.58—1.80 © mm (3), wingspan 3.8—4.1 mm. Antennae with 23—25 segments (3). Underside forewing and hindwing without special scales. Male genitalia (figs. 107, 250, 298, 369, 410). Capsule very short, length 150—167 um (3). Tegumen produced into ventral globular pseu- duncus (fig. 410). Gnathos (fig. 298) with cen- tral element broad and truncate, in form of a transverse bar. Valva (fig. 250) length 133—150 um, basally broad, below middle suddenly nar- rowed and inner margin becoming concave to- wards tip; outer margin uniformly convex. Ae- deagus (fig. 369) 244—279 um (3), more than 1.5 X as long as capsule, carinae single, pointed. Female genitalia (figs. 176, 439). T7 without long setae. T8 small, with few scales laterally and with 8—14 setae. Anal papillae with 6—8 setae. Vestibulum with vaginal sclerite, slightly different from that in other species and a spicu- late pouch with very few small spines, without pectinations. Corpus bursae small, 310—350 um, covered with many pectinations, except distal part; signa similar, oval, 189—223 um (209.3 + 14.9, 6), 2.0—2.3 X as long as wide. Ductus spermathecae with 3—3¥, convolutions. Larva emerald green, head-capsule brown. No ventral plates (Klimesch, 1978). Biology. Hostplant: Quercus macrolepis Kotschy. Mine (fig. 482). Egg on leaf upperside. Early mine contorted gallery, widening into irregular blotch or wide gallery, with dispersed central frass. Life history. Probably univoltine. Larvae col- lected in September, adults emerged in April. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 43 Distribution (fig. 546). Only known from Rhodos. Material examined: 3 d, 3 © (holo- and paratypes), ' Greece, Rhodos, Rodini, e.l. 17—30.iv.1973, Quercus macrolepis, 22.ix.1972, J. Klimesch (ZSMK). Mines: 2 mines, same data (ZMA). The Ectoedemia subbimaculella group This is a uniform group of Quercus mining species, making gallery mines or gallery-blotch mines. Adults of this group have various colour pat- terns, but never with metallic shining spots or fasciae. Males of most species possess costal bristles instead of a hair-pencil, except quın- quella, cf. algeriensis and gilvipennella. The group is best characterised by the female | genitalia: vestibulum with a ring-shaped vaginal sclerite, a spiculate pouch with the spicules partly separate, partly in small rows of 2—3 in contrast to populella-group, and a patch of dense pectinations near entrance of ductus sper- mathecae. In contrast with all other Ectoedemia species except intimella, the corpus bursae is de- L void of pectinations.The signa are long and elongate, dissimilar, the shortest being 3.5—7.5 times as long as wide, except in leucothorax. Larvae are yellow, whitish or green, and many species possess black ventral plates during the penultimate instars. The group is best developed in the mediterra- nean area, and also occurs in Japan. 20. Ectoedemia (Ectoedemia) quinquella (Bedell, 1848) He 106177251299 570) Al L440) 485° 527) Microsetia quinquella Bedell, 1848: 1986. Syntypes, England, West Wickham, 30.vi.1847, G. Bedell, (depository unknown), [not examined] [no genus] quinquella; Herrich-Schäffer [1854]: pl. 114 fig. 928. Nepticula quinquella; Stainton, 1849: 29; 1854: 301; Herrich-Schaffer, 1855: 355; Frey, 1857: 407, 408; Stainton, 1859: 433; Wocke, 1871: 339; Mey- mak, 18775 iil, UWS 19958 7259 gi WEE Sr 343; Rebel, 1901: 227; Meess, 1910: 480; Mey- rick, 1928: 862; Petersen, 1930: 76, fig. 114 (d genitalia). Dechtiria quinquella; Beirne, 1945: 206, fig. 71 (d genitalia); Emmet, 1971: 248. Stigmella quinquella; Gerasimov, Lhomme, 1963: 1201. Stigmella (Dechtiria) quinquella; Hering, 1957: 870, fig. 534 (mine). Trifurcula (Ectoedemia) quinquella; Johansson, 1971: 245. 1952 2552 Ectoedemia quinquella; Bradley et al., 1972: 2; Em- met, 1976: 189, pl. 6, fig. 16, pl. 12, fig. 33. Diagnosis: easily separated from all other Ectoedemia species, described here, except alge- riensis, by characteristic pattern of three white spots on forewing: a costal, a dorsal and a discal spot. It can be separated from algeriensis by its dark thorax, and males from cf algeriensis by the darker hair-pencil and different form of val- va and gnathos. Description. Male. Forewing length 1.84—2.28 mm (2.10 + 0.17, 6), wingspan 4.2—5.0 mm. Head: fron- tal tuft almost completely black, with a few fus- cous scales on frons; collar black. Antennae with 36—42 segments (39.8 + 2.6, 4). Thorax black, posterior tips of mesoscutum and tegulae white. Forewings black with three white spots: a costal on Vs from wingbase, a dorsal, approxi- mately in middle, and a discal on % from base, sometimes a few white scales near wingbase. Hindwing with yellowish hair-pencil of approx- imately Va hindwing length, surrounded by yel- low lamellar scales. Female (fig. 55). Forewing length 2.04—2.68 mm (2.37 + 0.19, 8), wingspan 4.6—5.6 mm. Antennal segments 26—29 (28.1 + 1.1, 7). Male genitalia (figs. 108, 251, 299, 370, 411). Capsule length 227—266 um (2). Tegumen (fig. 411) rounded, slightly indented at tip. Gnathos (fig. 299) with central element divided, distal part spatulate, basal part with serrate margin. Valva (fig. 251) length 171—257 um (2), inner margin concave, except basally, tip narrow, dorsal surface with comparatively few setae. Aedeagus (fig. 370) length 171—257 um (3), ca- rinae pointed, single or bifurcate, sometimes with additional spines at base. Female genitalia (figs. 177, 440). T8 with two lateral groups of scales and few setae, on T7 along anterior margin of T8 a few small setae, not arranged in distinct row. Anal papillae with 12—18 setae. Vestibulum with vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectinations near entrance of ductus spermathecae. Corpus bursae 550—670 um, without pectinations; signa dissimilar, longest 411—514 um (4), shortest 356—454 um, 4.0— 4.7 X as long as wide (4). Ductus spermathecae with 2 indistinct convolutions. Larva. Yellow, with dark brown head-capsule and conspicuous black ventral plates, which are shed during final instar. Thereafter ganglia visi- ble. 44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Biology. Hostplants: Quercus robur L. and Q. petraea el: Mine (fig. 485). Egg on leat underside, often against vein. Mine highly contorted gallery; ear- ly mine filled with narrow linear frass, later with irregular dispersed black frass, leaving wide clear margins. Often many mines occur in the same leaf. -Life history. Univoltine. Larvae occur late in the season, in England in late October and No- vember, in Greece very young larvae have been found in mid September. The adults fly in the second half of June and early July. Distribution (fig. 527). Atlantic-mediterranean species, locally abun- dant in southern England, known from a small number of localities in Belgium, France, Italy and Greece. Record from Norway (Grönlien, 1937) probably incorrect. Remarks. Types of this species are unknown, and Be- dell’s collection does not seem to exist any more (see atrifrontella). From Bedell’s description and figure the identity of this species is not in doubt. Material examined: 6 d, 119. — Belgium: 1 9, Tervuren, 20.vi.1888, Crombrugghe; 1 ©, Zolder, 27.v1.1938, E. Janmoulle (IRSN). — France: 1 ®, Achères (Yvelines), 22.vi.1947, Le Marchand; 1 6, l’Etang la Ville (Yvelines), 21.vi.1942, Le Marchand (MNHN); 1 6, Vannes, e.l. 27.vi.1913, Joannis (IRSN). — Great Britain: 2 d, 19, 10 km NE New- market, Herringswell, 11.x1.1981, e.l. 8—10.vi.1982, A. M. Emmet, J. W. Schoorl; 1 2, Pods Wood, 2 km N. of Tiptree (Essex), 23.x.1979, e.l. 17.vi.1980, A. M. Emmet, G. Bryan & E. J. van Nieukerken; 2 6, 4 2,3 km E. Rainham, Belhus Wood, 24.x.1979, e.l. vi.1980, G. Bryan, E. J. van Nieukerken (ZMA, partly on al- cohol). — Greece: 1 2, Litochorion, 3—400 m, 14— 22.v1.1957, J. Klimesch (ZSMK). — Country un- known: 1 ©, Macedonia, Kr., coll. Staudinger (MHUB). Mines. — On Quercus robur. — Great Britain: Herringswell; Tiptree; Rainham; Weeley. On Quer- cus petraea sl. — Greece: 4 km W. Palaiokastron, Evritania. Additional records (figs. of externals and d genita- lia by Klimesch, examined). — Italy: Liguria, Testico (near Alassio), 470 m, 5.vii.1969, Jackh; Liguria, Con- na, S. Sebastiano (near Pigna), 4.vii.1969, Jackh. 21. Ectoedemia (Ectoedemia) algeriensis sp.n. (figs. 56, 178, 441, 484, 527) Type material: Holotype 9: Algeria: Aures, near Arris, 32 km SSE of Batna, 1700 m, 28 1V 1280 MopentO Wes wes, Sit 25, El 13.vi.1980, Quercus ilex, VU no 80064 KE, Bryan, van Nieukerken & Oosterbroek, Geni- talia slide 1125 (ZMA). Paratypes, 2 2, same data as holotype, e.l. 13—16.v1.1980 (BMNH, ZMA); Mines examined from type locality and from Algeria: Aures, Dj. Chélia, 1600—1900 m. Diagnosis: externally very similar to quin- quella, but thorax entirely white and basal white spot present. Genitalia (2) very characteristic by dense hairy abdominal tip. Description. Male. Unknown, but see below. Female (fig. 56). Forewing length 2.28—2.56 mm (3), wingspan 5.0—5.6 mm. Head: frontal - tuft and collar fuscous to black. Antennae with 27—33 segments (3). Thorax completely white. Forewings black, with four white spots: a small basal, a large costal before middle, a dorsal, ap- proximately in middle and a discal at 24 from wingbase. Female genitalia (figs. 178, 441). T8 (and T7?) with more than 70 long setae, partly in row along anterior margin, no scales. Anal papillae with 24—28 setae. Vestibulum with vaginal sclerite, a prominent dorsal spiculate pouch, and a group of densely packed pectinations near en- trance of ductus spermathecae. Corpus bursae 605—660 um without pectinations; signa dissi- milar, longest 386—450 um (2), shortest 355— 420 um, 3.5—3.9 X as long as wide (2). Ductus spermathecae with 2 indistinct convolutions. Larva. Green, without ventral plates. Not ex- amined in detail. Biology. Hostplant: Quercus rotundifolia Lam. (often regarded as form of ilex). Mine (fig. 484). Egg on upper surface, often on or near vein. Gallery, much contorted with black frass leaving narrow clear margins. Mine similar to that of ilicis, heringella and haraldi, only separable by colour of larva. Life history. Larvae taken in late April, adults emerged in June. Males of cf. algeriensis found in July. Distribution (fig. 527). Algeria: Aurés mountains, and probably Mo- rocco (see remarks). Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 45 Remarks. This species is described from three females reared from a small sample, from which unfor- | tunately no males emerged. Although clearly re- | lated to quinquella, it is a distinct species, differ- ing in genitalia and biology. The males de- | scribed below probably belong to algeriensis because, although they resemble quinquella, they also differ in some ways. It is however not wise to include them in the type-series of alge- riensis, since they are too worn. Also they have not been reared. I have reared 1 ®, (slide 1897) from Quercus coccifera from Spain (Malaga): 7 km N. Benaha- vis, road to Ronda, 800 m, 7.11.1984, e.l. 17— 18.iv.1984, which externally corresponds with | algeriensis, and also in the internal genitalia. However, the terminal segments differ (fig. 442) from those of the type series, and I therefore can not identify this specimen with certainty * until further material is available. 21A. Ectoedemia (Ectoedemia) cf. algeriensis sp.n. (male) (figs. 109. 252, 300, 371) Material: 2 &, Morocco: Moyenne Atlas, Az- rou, 16.vu.1975, F. Kasy (NMW). Two worn males, which probably belong to algeriensis, see remarks on that species. Diagnosis: wing pattern unknown, differs from ilicis and heringella by presence of hair- pencil, from quinquella by white hair-pencil, and from all by large number of antennal seg- ments. Genitalia similar to ilicis and heringella, but central element of gnathos remarkably large. Description. Male. Forewing length 2.4 mm, wingspan + 5.4 mm. Head: colour of frontal tuft unknown, all scales lost in the two specimens. Antennae long, with 53—54 segments. Thorax probably white. Colour-pattern of forewing not recogni- sable, but presence of discal spot likely, the dis- tribution of the few scales left on the wings, suggest the likelyhood of a similar pattern as al- geriensis. Hindwing with a white hair-pencil, surrounded by a patch of yellow scales. Male genitalia (figs. 109, 252, 300, 371). Cap- sule 257 um long. Tegumen rounded. Gnathos (fig. 300) with central element divided, distal part prominent, spatulate, basal part with ser- rate margin. Valva (fig. 252) length 206 um, in- ner margin concave, tip wide and truncate, dor- sal surface with few setae. Aedeagus (fig. 371) 274 um, carinae pointed, bi- or trifurcate. 22. Ectoedemia (Ectoedemia) gilvipennella (Klimesch, 1946) comb. n. (figs. 57, 58, 110, 179; 253, 301, 372, 443, 486, 543) Stigmella gilvipennella Klimesch, 1946: 168, fig. 8. Lectotype ¢ (here designated), Italy: Liguria, Fer- rania near Altare, el. 26.1v.—7.v.1945, Quercus cerris, ix.1944, Zucht No. 509, J. Klimesch, Geni- talia slide Kl. 272 (ZSMK) [examined]. Stigmella (Stigmella) gilvipennella; Hering, 1957: 870. Nepticula (Stigmella) gilvipennella; Szöcs, 1968: 228. Diagnosis: the only predominantly white Ectoedemia, further characterised in the male by the prominent fuscous or black hair-pencil. The other uniformly coloured Ectoedemia species are darker and often larger. Without examining genitalia or venation, females could be mistaken for Trifurcula or Acalyptris species. Male genitalia very similar to those of quin- quella, but separated by dorsal lobe of valva. Description. Male (figs. 57, 58). Forewing length 2.08— 2.48 mm (2.32 + 0.13, 12), wingspan 4.9—5.4 mm. Head: frontal tuft yellowish, mixed with fuscous, especially on vertex; collar yellowish white. Antennae with 28—34 segments (30.8 + 1.5, 11). Thorax and forewings predominantly white, irrorate with dark brown tipped scales, no distinct colour-pattern. Hindwing with fuscous to black hair-pencil of % hindwing length, not surrounded by special scales. Female. Forewing length 1.96—2.36 mm (2.21 + 0.13, 13), wingspan 4.4—5.2 mm. An- tennal segments (17)23—24 (23.4 + 0.5, 10). Male genitalia (figs. 110, 253, 301, 372). Cap- sule length 210—240 um (219.4 + 11.9, 5). Te- gumen rounded. Gnathos (fig. 301) with central element divided, distal part spatulate, basal part with serrate margin. Valva (fig. 253) length 171—193 um (177 + 8.9, 5), inner margin con- cave, outer margin dorsally folded back, form- ing an inwardly projecting lobe, covering sever- al setae, tip pointed. Aedeagus (fig. 372) 244— 257 um (248.6 + 6.1, 4), carinae pointed, single. Female genitalia (figs. 179, 443). T7 with a row of 8 long setae along anterior margin of T8; T8 with 8 setae, no scales. Anal papillae with 11—13 setae. Vestibulum with vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectinations near entrance of ductus spermathecae. Corpus bursae 500 um, without 46 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 pectinations; signa dissimilar, longest 347 um (1), shortest 330 um (1), 3.9 X as long as wide. Ductus spermathecae with 3 convolutions. Larva. Bright emerald green, with light yel- low head-capsule. Ganglia invisible, ventral plates absent. Biology. Hostplant: Quercus cerris L. „Mine (fig. 486). Egg on leaf upperside, often on or near vein. Early mine: narrow contorted gallery with broken brown frass, later becoming wide and more contorted gallery filled with brown dispersed frass. Life history. Univoltine. Larvae from late October until late November. Adults reared from April to June. Distribution (fig. 543). Probably throughout the range of Quercus cerris, but yet only recorded from northwest Italy, Hungary, and here for the first time from Austria and Yugoslavia. Material examined: 14 6, 13 2. — Austria: 6 6, 8 2, Hof am Leithagebirge, S. of Mannersdorf (Nie- deröst), 200 m, el. 30.iv—14.vi.1984, E. J. van Nieu- kerken; 2 5, 3 2, Loretto, 7 km N. Eisenstadt (Bur- genland), 240 m, el. 30.iv.—8.v.1984, E. J. van Nieu- kerken (ZMA). — Hungary: 1 d, Törökbálint (W. of Budapest), 27.1v.1965, e.l., J. Szöcs; 3 d, 2 ®, same data, el. 13—24.v.1974 (TMAB). — Italy: 2 d (lecto- and paralectotype), Liguria, Ferrania near Altare, e.l. 26.iv.—7.v.1945, J. Klimesch (ZSMK). Mines. — Austria: Hof am Leithagebirge; Loretto. — Hungary: Törökbálint. — Yugoslavia (Bosna): S. of Han Knezica, 11 km N. of Prijedor. 23. Ectoedemia (Ectoedemia) leucothorax sp. n. (figs. 59, 111, 180, 181, 254, 302, 372, 444, 527) Type material: Holotype 6, Spain, Marbella (Malaga), 5.v.1981, C. Gielis, Genitalia slide VU 1892 (ZMA). Paratypes, 2 6,3 2. — Spain: 1 2, Andalusia, Marbella, L. Monteros, 25 m, 12.vii.1972, E, Traugott-Olsen (ZMC); 1 di, Andalucia, Camino de (road to) Ojen, 150 m (Marbella), 25.vi.1983, E. Traugott- Olsen (ETO); 1 g, 2 2, Estepona, 10-21.v1.1979, Leo Kohonen (ZMUO, ZMA). Diagnosis: easily recognised by white thorax, orange head and forewing with white streak along dorsal margin, running from base to fas- cia, and in male absence of hair-pencil. Exter- nally most sımilar caradjaı has a dark thorax and hair-pencil. Male genitalia characterised by very long, slender valvae and aedeagus shorter than capsule or valvae; female genitalia by widened anterior apophyses, shape of T8, hairy abdominal tip, similar signa and smooth bursa. Description. Male. Forewing length 2.28—2.44 mm (2), wingspan 5.2—6.0 mm. Head: frontal tuft and collar intensively orange. Antennae with 41—42 segments (2). Thorax and tegulae white, except brown outer edge of tegulae; forewings fuscous, with medial arched or interrupted white fascia, united by white streak along dorsal margin to wingbase, occupying 3—4 rows of scales; white pattern in rest position of moth forming anchor- shaped figure. Hindwing without hair-pencil, but with costal bristles. Female (fig. 59). Forewing length 2.4—2.72 | mm (3), wingspan 5.2—6.0 mm. Antennae with 31—32 segments (3). Male genitalia (figs. 111, 254, 302, 372). Cap- sule length 304—330 um (3). Tegumen pro- duced into rounded, approximately triangular, pseuduncus. Gnathos (fig. 302), divided, with narrow spatulate distal part, basal part with ser- rate margin. Valva (fig. 254) length 279—321 um, very long and narrow, inner margin com- pletely concave, outer margin completely con- vex. Aedeagus (fig. 372) 244—279 um (3), dis- tinctly shorter than capsule or valva, with single pointed carinae, curved outwards. Female genitalia (figs. 180, 181, 444). T7 with a semicircular patch with about 200 closely set long, smooth setae. T7 and 8 in addition with about 50 shorter setae and a few scales laterally; T8 with posteror margin truncate with promi- nent corners. Anal papillae broad, with 16 setae. Vestibulum with vaginal sclerite, a dorsal spicu- late pouch with many small spicules and a group of densely packed pectinations near entrance of ductus spermathecae. Corpus bursae 620—660 um, without pectinations; signa similar, 309— 339 um, 2.6—3.4 X as long as wide. Ductus spermathecae with 2 convolutions and a promi- nent vesicle. Larva unknown. Biology. Hostplant unknown, but most likely ever- green Quercus, judging from its relationships and localities. In the Marbella localities Quercus suber or Q. coccifera grow. In February 1984 I was not able to collect there any other mines Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 47 | than those similar to E. suberis, but it is possible | that leucothorax feeds in another season. Life history. Adults found from early May to early July, all collected at light. Distribution (fig. 527). Only known from the Costa del Sol in Spain. Remarks. This species shows some similarities with the | suberis group, but the absence of pectinations in the bursa, the presence of a group of pectina- tions in the vestibulum, the form of the gnathos and the presence of costal bristles in the male in- dicate that it in fact belongs to the subbimaculel- la group. The presence of many long setae on the female abdominal tip probably is an adapta- | tion to oviposition on rough surfaces of ever- green oak leaves, and hence a parallel devel- opment with suberis and algeriensis. 24. Ectoedemia (Ectoedemia) haraldi, (Soffner, 1942) (figs. 60, 112, 182, 255, 303, 374, 445, 487, 545) Nepticula haraldı Soffner, 1942: 56, figs. 1—12. Lec- totype d (here designated), France: Angouleme, el. v.1941, Quercus ilex, 11.1941, Zucht No. 382a, Soffner, Genitalia slide 4776 (MHUB) [examined]. | Stigmella prinophyllella Le Marchand, 1946: 285. Ho- | Ectoedemia (Dechtiria) haraldi; Klimesch, lotype © [in description as 4], France: Villenave d’Ornon, Gironde, e.l. 23.v.1928, Le Marchand, Genitalia slide VU 0941 (MNHN) [examined]. (Synonymised by Le Marchand, 1948). Stigmella haraldı; Hering, 1957: 867, fig. 553 (mine); Lhomme, 1963: 1196. 1975a: 864, figs. 5, 6(¢ genitalia). Trifurcula (Ectoedemia) haraldi; Leraut, 1980: 49. Nepticula ilicella Constant [nomen nudum]. (Synony- mised by Klimesch, 1975a: 864. Diagnosis: externally very similar to albifas- ciella complex and preisseckeri, but with gener- ally lighter appearance. E. ilicis and heringella can be separated by the absence of a costal spot, and androconial scales in male heringella. E. su- beris can be distinguished by the straighter fas- cia and by the presence of a hair-pencil in male and hairy abdomen tip in female. Females of an- dalusiae are very similar to haraldi, and can on- ly be identified with certainty by genitalia. Male genitalia very characteristic by shape of valva with bulgy outer margin. Female genitalia char- acterised by wide T8 and wide, rounded S8. Description. Male. Forewing length 2.88—3.32 mm (3.07 + 0.13, 8), wingspan 6.2—7.1 mm. Head: fron- tal tuft light yellow to yellowish orange; collar similar. Antennae with 35—42 segments (37.8 + 2.4, 8). Thorax brown, sometimes mesoscu- tum with white tip. Forewings brown, with a white dorsal spot in middle, and a costal spot before middle, sometimes united to form a fas- cia. Hindwing without hair-pencil, but with costal bristles. Female (fig. 60). Forewing length 2.56—2.88 m (2.75 + 0.13, 10), wingspan 5.8—6.5 mm. Antennal segments 27—31 (29.1 + 1.5, 8). Fe- male distinctly smaller than male. Male genitalia (figs. 112, 255, 303, 374). Cap- sule length 266—300 um (286.3 + 14.7, 5). Te- gumen rounded. Gnathos (fig. 303) with central element divided, distal part truncate, basal part with serrate margin. Valva (fig. 255) length 193—206 um (201.4 + 5.2, 5), outer margin bulging distally, inner margin basally straight or convex, from 1/3 distinctly concave, up pro- nounced, pointed. Aedeagus (fig. 374) 274—283 um (279.4 + 3.6, 5), carinae varying from single to multifurcate. Female genitalia (figs. 182, 445). T7 with only few short setae along anterior margin of T8, not in distinct rows. T8 with two lateral groups of scales and 3—5 setae each; posterior margin al- most straight, lateral corners pronounced, rounded; S8 broadly rounded. Anal papillae with 14—23 setae. Vestibulum with vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectinations near the entrance of ductus spermathecae. Corpus bursae 570—825 um, without pectinations; signa dissimilar, longest 363—577 um (460 + 56, 11), shortest 308—495 um (402 + 48, 11), 4.05.4 X as long as wide. Ductus spermathecae with 2 indistinct convolutions. Larva. Whitish, opaque, with distinct brown ganglia. Head-capsule and prothoracic plate dark brown. Ventral plates absent. Biology. Hostplants: Quercus ilex L., Q. rotundifolia Lam. and Q. coccifera L. Not yet recorded from Q. suber L., but probably also feeds on that species. Mine (fig. 487). Egg on leaf upperside, not against vein. Early mine: slightly contorted nar- row gallery, gradually widening, remaining lin- ear throughout. Filled with thick black frass, hardly leaving clear margins. Not always sepa- rable from mines of algeriensis, ilicis or heringel- la. 48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Life history. Univoltine. Larvae collected in February and March, adults from April to June. Distribution (fig. 545). Widespread in southern France, occurring along Atlantic coast up to Angouleme, further recorded from Spain, Portugal, Italy and Greece. _ Remarks. Syntypes of haraldi are present in many col- lections. A lectotype is here designated from the Hering collection in Berlin, since it contains a large number of nepticulid types. The Soffner collection is not housed there. Marchand incorrectly gave the holotype of prinophylella as male. He was the first to sepa- rate this species from suberis, with which it had been confused earlier. Material examined: 18 6, 26 2. — France: 4 4,49 (lecto- and paralectotypes of haraldi), Angouleme, e.l. v.1941, Quercus ilex, J. Soffner (MHUB, ZMA, ZMC); 1 9, Bize, v.1909, Chrétien; 2 d, 2 2, Alpes marit., Cannes, 13, 15 [decade], Constant (MNHN); 2 6,1 2, Golfe Juan, Alpes maritimes, 8—15.vi.1894, Constant (BMNH); 1 d, “Nesp.” (? near St. Pons, dep. Hérault), 15.vi.1904, Chrétien; 2 d, 2 2, Roque- fort (B. du Rh.), between Cassis and Cuges les Pins, el. 14—24.1v.1984, Quercus ilex, R. Buvat (ZMA); 1 2, Viens (Vaucluse) (near Apt), el. 5.v.1971, Quercus ilex, R. Buvat (coll. Buvat); 3 d, 1 2 (holo- and para- types of prinophylella), Villenave d’Ornon, Gironde, el. 23.v.—1.vi.1928, Quercus ilex, Le Marchand (MNHN). — Greece: 1 ©, Lakonia, 7 km SW Mo- nemvasia, 9.iv.1981, B. Skule (ZMC). — Italy: 3 9, Sistiana Mare, 0—60 m, e.l. 6—9.v.1970, Quercus ilex, G. Deschka (LNK). — Portugal: 2 9, San Fiel, el. 20.iv, Quercus coccifera, [Mendes], coll. Joannis; 3 d, 4 9, [prov. Beira Baixa, San Fiel] e.l. 23.iv, Q. dex, [Mendes], coll. Joannis; 1 d, (misidentified paralecto- type of cs Mendes) idem, el. 22.v. (MNHN). — Spain: 3 9, 7 km N. Benahavis (Málaga), road to Ron- da, 800 m, el. 3—18.iv.1984, Quercus coccifera, E. J. van Nieukerken (ZMA). Mines. — On Quercus coccifera. — Spain: 7 km N. Benahavis. On Quercus ilex. — France: Angoulème, leg. Soffner (BMNH); between Cassis and Cuges les Pins, leg. Buvat. 25. Ectoedemia (Ectoedemia) ilicis (Mendes, 1910) comb. n. (figs. 61, 113, 183, 256, 257, 304, 375, 446, 488, 489, 543) Nepticula ilicis Mendes, 1910: 164, pl. 7 figs. 7, 8. Lec- totype d (here designated), Portugal: [San Fiel, prov. Beira Baixa] e.l. 22.v., Q. ilex, [Mendes], Chenille a téte noire, Coll. L. & J. de Joannis, Genitalia slide VU 1358 (MNHN) [examined]. Stigmella ilicis; Gerasimov, 1952: 243; Hering, 1957: 869 (mine). Diagnosis: ilicis and heringella are the only western Palaearctic oak-mining species with dorsal spot only. Fomoria septembrella (Stain- ton), Stigmella catharticella (Stainton) and Zim- mermannia species also have dorsal spot only, but this is situated postmedially, whereas it is medial in zlıcıs. This is also the case in E. inti- mella, but this species can be separated by its ‘unicolorous antennae, uniform dark scales on the forewings, and hair-pencil in the male. See heringella for differences with that species. The mines are easily confused with haraldi, but adults are easily separated by totally different valva in male and the distinct row of setae on T7 and form of T8 in female of ilzcis. Description. Male (fig. 61). Forewing length 2.48—3.36 mm (2.87 + 0.25, 13), wingspan 5.6—7.2 mm. Head: frontal tuft and collar yellowish orange. Antennae with 31—40 segments (37 + 2.5, 13); scape white, with sometimes some brown scales. Thorax and forewings brown, with a dorsal spot only in medial position, sometimes slightly extending along dorsal margin towards base; sometimes a few scattered white scales present in addition. Hindwing without hair- pencil, but with costal bristles. Female. Forewing length 2.36—2.88 mm (2.68 + 0.16, 10), wingspan 5.1—6.5 mm. An- tennal segments 28—31 (30.1 + 1.0, 8). Male genitalia (figs. 113, 256, 257, 304, 375). Capsule length 231—244 um (240 + 6.1, 5). Te- gumen broad and rounded. Gnathos (fig. 304) with central element undivided, slightly trun- cate, lateral margins serrate. Valva (figs. 256, 257) length 176—193 um (183.4 + 7.0, 5), inner margin basally straight or convex, from 1/3 dis- tinctly concave, inwards pointed tip prominent, truncate. Aedeagus (fig. 375) 253—274 um (264 + 9.9, 5), carinae split into two or more spines each. Female genitalia (figs. 183, 446). T7 with a distinct row of 8-14 long setae along anterior margin of T8. T8 with two groups of about 3— 6 setae, scales absent; T8 narrow with slightly sinuous posterior margin. Anal papillae with 8- 14 setae. Vestibulum with vaginal sclerite, a dorsal spiculate pouch and a group of densely packed pectinations near the entrance of ductus spermathecae. Corpus bursae 660—825 um, VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 49 without pectinations; signa dissimilar, longest 407—471 um (432.9 + 26.9, 5), shortest 369— 416 um (395.1 + 19.3, 5), 3.9—5.1 X as long as wide. Ductus spermathecae with 2 indistinct convolutions. Larva. Yellow with conspicuous brown gan- glia. Head light brown. Ventral plates absent. Biology. Hostplants: Quercus ilex L., Q. rotundifolia Lam. and Q. suber L. Often sympatric with ha- raldi and suberis. Mine (fig. 488, 489). Egg on leaf upperside, usually against vein. Early mine: much con- torted gallery, starting very narrow. Frass black, dispersed, leaving narrow clear margins. Mine seems longer and more contorted than in haraldı, but difficult to separate. Life history. Univoltine. Larvae found in Jan- uary and February. Adults from March to the end of June. Distribution (fig. 543). Clearly west mediterranean. Remarks. As in the case of Parafomoria ladaniphila (Mendes) (Van Nieukerken, 1983: 469) type material of ilicis seems no longer to exist in Por- tugal, but Portuguese material in de Joannis col- lection (MNHN) can be regarded as syntype material if labelled as ilicis. I examined 3 d and 1 2 mounted on the same block of pith and la- belled “ilicis Mendes”. One of these males be- longs to haraldi, but the other specimens are this species. Since Mendes clearly refers in his description to the species with a dorsal spot on- ly, one male of the two is selected lectotype, and the haraldi male is regarded as a misidentified paralectotype. Later I found more paralecto- types in the Hering collection in Berlin. Material examined: 14 d, 12 9. — Algeria: 1 d, Batna, 1.v.1903, Walsingham (BMNH). — France: 1 2, Bize, 30.vi.1910, Chrétien; 1 6; 3 2, “Nesp.” (? near St. Pons, dep. Hérault), 15.vi.1904, Chrétien (MNHN). — Portugal: 3 5, 2 © (lecto- and paralec- totypes), [San Fiel, prov. Beira Baixa], e.l. 22+ 26.v, | Quercus ilex, [Mendes] (MNHN, MHUB). — Spain: 1 ®, Sierra de Alfacar (near Granada), 24.1v.1880, Staudinger (MHUB); 1 4, Marbella, El Mirandor, 100 m, 17.v.1969, E. Traugott-Olsen (ETO); 2 ®, Port Bou, el. 29—30.111.1968, Quercus ilex, J. Klimesch (ZSMK); 7 d, 3 2, 4 km NE. Igualeja, Serrania de Ronda (Malaga), 1100 m, el. 19.11—16.1v.1984, | Quercus rotundifolia, E. J. van Nieukerken; 1 4,1 ©, Sierra Blanca, 6 km N. Marbella (Malaga), El Mira- dor, 800 m, el. 17—24.iv.1984, Quercus rotundifolia (2)+ Q. suber (d), E. J. van Nieukerken (ZMA). Mines. — On Quercus rotundifolia. — Portugal: San Fiel, leg. Mendes (BMNH). — Spain: Serrania de Ronda; Sierra Blanca. On Quercus suber: Spain: Sier- ra Blanca. Additional record. — France: 1 6, 1 2, Marseille, el. 17.v.1971, 27.v.1972, Quercus ilex, R. Buvat (R. Johansson, pers. comm.). 26. Ectoedemia (Ectoedemia) heringella (Mariani, 1939) comb. n. (figs. 62—64, 114, 115, 185, 186, 258, 259, 305, 306, 376, 377, 447, 448, 544) Nepticula heringella Mariani, 1939: 5, 6, fig. 1a, pl. 1. Lectotype d (here designated), Italy: Sicilia, Parti- nico, 1.v.1937 [Quercus ilex], Mariani (MCST) [examined]. Nepticula heringella f. alliatae Mariani, 1939: 7. Stigmella heringella; Hering, 1957: 868, fig. 554 (mine). Diagnosis: very similar to zlıcıs, but male easi- ly separated (also from most other species) by patches of brown androconial scales on hind- wing upperside and forewing underside. Female cannot always be separated with certainty from has, but usually heringella has some white scales in the region of the costal spot and also has slightly longer signa. Description. Male (figs. 62, 63). Forewing length 2.08— 2.68 mm (2.43 + 0.13, 19), wingspan 4.4—6.0 mm. Head: frontal tuft yellowish white to orange, in specimens from Cyprus fuscous on vertex; collar yellowish white. Antennae with 35—42 segments (38.4 + 1.8, 15); scape with some brown scales in posterior distal corner. Thorax and forewings brown with some scat- tered white scales; medial dorsal spot white, some white scales along costa, not forming a distinct costal spot; underside of forewings with basally an elongate patch of brown (androconi- al) scales. Hindwing without hair-pencil, but with costal bristles; in basal half with a patch of brown (androconial) scales on upperside. Female (fig. 64). Forewing length 2.24—2.60 mm (2.44 + 0.20, 14), wingspan 4.6—5.8 mm. Antennal segments 27—32 (29.9 + 1.4, 16). Without patches of brown scales on underside forewing or upperside hindwing. Male genitalia (figs. 114, 115, 258, 259, 305, 306, 376, 377). Capsule length 236—283 um (252.9 + 16.5, 9). Tegumen broad and rounded. 50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Gnathos (figs. 305, 306) with central element di- vided, distal part spatulate, basal part with ser- rate margin. Valva (figs. 258, 259) length 180— 223 um (209.0 + 17.2, 9), inner margin almost straight or concave, tip prominent, slightly truncate. Aedeagus (figs. 376, 377) 257—300 um (274.3 + 16.5, 9) carinae single, bi- or trifur- cate. Female genitalia (figs. 185, 186, 447, 448). T7 with a distinct row of 8—12 long setae along anterior margin of T8. T8 with two groups of 2—4 setae (7 in Cyprus specimen), without scales, but some scales present in specimen from Corsica; T8 narrow, with slightly sinuous pos- terior margin. Anal papillae with 8—15 setae. Vestibulum with vaginal sclerite, a dorsal spicu- late pouch and a group of densely packed pecti- nations near the entrance of ductus spermathe- cae. Corpus bursae 580—715 um, without pec- tinations; signa dissimilar, longest 407—583 um (484 +73, 5), shortest 353—517 um, (116 + 73, 5), 4.04.7 X as long as wide. Ductus sperma- thecae with 2 indistinct convolutions. Larva not examined. Biology. Hostplants: Quercus ilex L., Q. alnifolia Poech (on Cyprus). Mine. Egg on leaf upperside, often near vein. Mine: much contorted gallery, amost filled with black frass. Not to be separated from mine of ilicis. Life history. Univoltine. Larvae taken from November to April (Hering, 1957). Adults from late April to the end of June. Distribution (fig. 544). From Corsica eastwards to Cyprus. Seems to be the eastern vicariant of E. ilicis. Not yet re- corded from Greece. Remarks. This species shows some variability. The specimens from Cyprus differ in darker head- colour and some genitalic details, but, since they also have the diagnostic features of heringella, I regard these as conspecific with heringella. The form alliatae, described by Mariani has no taxo- nomic value, it is probably described from worn specimens. Material examined: 28 d, 26 9. — Cyprus: 1 à, 1 ©, Arakapos (Troödos mountains), el. 25.11.1980, Quercus alnifolia, B. Gustafsson (RMS). — France: 1 3, 1 2, Corsica: Corte, 14.vi.1899, Walsingham (BMNH). — Italy: 4 6, 3 9, Latina, Monti Aurunci, 5 km. N. Itri, 600 m, 24—30.vi.1969, R. Johansson (coll. Johansson); 2 2 (paralectotypes), Sicilia, Paler- mo, 8.vi.1928, Mariani; 1 6, 1 2, idem, el. Sileviel9375 Mariani (MEST) Crhiden sel 16.vi.1964, W. Glaser (LNK); 5 6, 6 ® (lecto- and paralectotypes), Sicilia, Partinico, 1—11.v.1937, Mari- ani (MCST, MHUB, ZMC). — Yugoslavia: 12 d, 18 2, Rijeka, Istria, 100 m, el. 6—20.v.1970, Quercus 1l- ex, G. Deschka (LNK); 1 6, Split, Dalmatia, 19.v.1959, Novak (TMAB); 3 6, 4 2, Zadar, Dalma- tia, 0-60 m, el. 13—24.v.1970, Quercus ilex, G. Deschka (LNK). Mines. — On Quercus alnifolia. — Cyprus: Araka- pos, leg. Gustafsson (RMS). On Quercus ilex. —Italy: Sicilia, Taormina, leg. Groschke (BMNH). 27. Ectoedemia (Ectoedemia) alnifoliae sp. n. (figs. 65, 187, 188, 449, 546) Trifurcula (Ectoedemia) sp.; Gustafsson, 1981b: 468, _ fig. 9. Type material: Holotype 2, Cyprus: Tro- ödos, 10.11.1979, [e.l. 17.1v.1979], Quercus alni- folia, [B. Gustafsson], Genitalia slide RMS 6572 (RMS). Mine from which holotype emerged ex- amined. Diagnosis: externally similar to nigrosparsel- la, but light scales not intensively yellow, and scape with scattered brown scales. Female geni- talia without long spiraled ductus spermathecae, with only 3 narrow convolutions. Description. Male unknown. Female holotype (fig. 65). Forewing length 2.88 mm, wingspan 6.6 mm. Head: frontal tuft orange, darker on vertex, collar yellowish. An- tennae broken, scape white with some brown scales. Thorax and forewings dark brown, irro- rate with some yellowish-white scales, no col- our pattern present. Female genitalia (figs. 187, 188, 449). T7 without a row of setae. T8 with two lateral groups of scales and approximately 5 setae each. Anal papillae with 15—18 setae. Vestibulum with vaginal sclerite, a prominent dorsal spicu- late pouch, and a group of densely packed pecti- nations near entrance of ductus spermathecae. Corpus bursae 690 um, without pectinations; signa dissimilar, longest 540 um, shorter 440 um, 4.0 X as long as wide. Ductus spermathecae with 2 narrow convolutions. Larva not examined. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 51 Biology. Hostplant: Quercus alnifolia Poech, an ever- | green oak. Mine. Egg on leaf underside. Mine starting as narrow gallery, suddenly enlarging into large blotch against leaf margin, frass not visible in single mine examined. See also Gustafsson (1981b: 469, fig. 9C). Life history. Larva taken in March, adult | emerged in April. Distribution (fig. 546). Troödos mountains on Cyprus. Remarks. Although only one female was available, this species is here described as new, since it shows sufficient diagnostic characters to separate it from other species, and the identity of males of this species can easily be determined by host- plant, mine-form and locality. 28. Ectoedemia (Ectoemedia) nigrosparsella (Klimesch, 1940) (figs. 66, 116, 189, 190, 260, 307, 378, 450, 491, 546) Nepticula nigrosparsella Klimesch, 1940a: 91, pl. 14 figs. 8, 9, pl. 15, figs. 10—12. Lectotype d (here designated). Italy: Teriolis merid., Naturno, near Merano, e.l. iv.1939, J. Klimesch, Genitalia slide 449/39 Hering (MHUB) [examined]. Stigmella nigrosparsella; Klimesch, 1951: 64; Hering, 1957: 869, fig. 543.(mine); Klimesch, 1961: 763. Ectoedemia nigrosparsella; Kasy, 1983: 5. Diagnosis: characterised by brown irrorate with yellow forewings and absence of hair-pen- cil in male. Male genitalia not separable from al- bifasciella complex. Female genitalia character- ised by long spiraled ductus spermathecae, with 1312—14 convolutions, whereas contorta usual- ly has 10%—12 convolutions (except one Spee men). Description. Male. Forewing length 2.0—2.68 mm (2.43 + 0.19, 9), wingspan 4.3—6.0 mm. Head: frontal tuft ferruginous, sometimes mixed with fus- cous; collar similar. Antennae with 28—37 seg- ments (32.5 + 3.2, 6). Thorax and forewings brown irrorate with light yellow scales, being a more pronounced yellow than in most other species; colour pattern absent, but light scales predominant at tornus. Hindwing without hair- pencil, but with costal bristles. Female (fig. 66). Forewing length 2.72—2.88 mm (2.79 + 0.07, 4), wingspan 6.0—6.4 mm. Antennal segments 25—27 (2.8 + 1.0, 4). Male genitalia (figs. 116, 260, 307, 378). Cap- sule length 283—309 um, (3). Tegumen round- ed. Gnathos (fig. 307) with central element truncate, as cut off. Valva (fig. 260) length 206—223 um (4), inner margin strongly convex, except apically, serrate by prominent setal sock- ets, tip pointed; dorsal surface with many setae. Aedeagus (fig. 378) 279—287 um (4), carinae pointed, single. Female genitalia (figs. 189, 190, 450). T7 with a row of 8—12 setae along posterior margin; T8 with two lateral groups of scales and 2—4 setae each. Anal papillae with 14—17 setae. Vestibu- lum with vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectina- tions near entrance of ductus spermathecae. Corpus bursae 740—825 um, without pectina- tions; signa dissimilar, longest 485—695 um (3), shortest 450—458 um, 4.1—4.4 X as long as wide (3). Ductus spermathecae with very prom- inent spiralised inner canal, with 13/2—14 con- volutions. Larva. Yellow, with greenish tinge in young- er larvae, head-capsule brown. In penultimate instars with conspicuous brown ventral plates, which are shed during final instar; thereafter the ganglia become visible. Biology. Hostplants: Quercus pubescens Willd., occa- sionally on Q. petraea (Mattuschka) Liebl. (Kli- mesch, 1951). Mine (fig. 491). Egg on leaf underside, occa- sionally on upperside. Early mine highly con- torted, forming brown blot with irregularly ac- cumulated brown frass; later gallery less con- torted, with brown dispersed or coiled frass, leaving narrow clear margins. Mine confined to small area, often near leaf-margin. Life history. Univoltine, larvae occurring from mid October to November. Adults col- lected at light mid June, reared in April and May (forced). Distribution (fig. 546). Known from a limited number of localities in Czechoslovakia, Hungary, Austria, Italy and France. Usually occurs in exposed southern slopes on calcareous soil — the typical habitat for Q. pubescens. 115) Oy Is ©. Glaslauterriegel, — Austria: 2 6, 10.vi.1983, Be Material examined: Gumpoldskirchen, 52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Kasy (NMW); 4 6, 5 9, ibid., e.l. 25.1v.—2.v.1984, Quercus pubescens, E. J. van Nieukerken (ZMA); 1 3, Hundsheimer Berg, Porta Hungarica (near Hain- burg), 19.vi.1976, F. Kasy (NMW). — France: 2 2, Aubagne (Bouches du Rhone), e.l. 10—17.v.1977, Quercus pubescens, Buvat (coll. Buvat). — Hungary: 4 6, 2 2, Törökbálint (W. of Budapest), el. 10— 18.iv.1974, 16.v.1976, Q. pubescens, J. Szöcs (TMAB). — Italy: 1 d, 1 2 (lecto- and paralecto- type), Naturno near Merano, e.l. iv.1939, J. Klimesch (MHUB); 3 6, 3 ©, Trento, Sardagna, 500 m, el. iv.1946, Q. pubescens, J. Klimesch (MHUB, ZMA). Mines. — Austria: Gumpoldskirchen; Hainburg; Loretto; Wien, Leopoldsberg. — Italy: Naturno, leg. Klimesch; Trento, leg. Klimesch (BMNH). The Ectoedemia albifasciella complex This is a complex of four sibling species com- parable with the subbimaculella complex, but differing in so far that the species are well sepa- rable on the female genitalia, the number of convolutions of the ductus spermathecae being a good and constant character in this complex: albifasciella with 2Y,—2¥,, cerris 3'%—4 pubescivora 5—6 and contorta with 10%—12 (131%) convolutions. The externals and male genitalia do not provide any diagnostic charac- ters. The species seem to have a different food- plant choice: albifasciella on Quercus robur and Q. petraea, pubescivora and contorta on Q. pubescens and cerris on Q. cerris, on which only one exception is known. Only E. albifasciella is described fully, the other species only as far as they differ. 29. Ectoedemia (Ectoedemia) albifasciella (Heinemann, 1871) Gress O67 17, ION, ISA, 2618505 0979; 416, 451, 490, 522) Nepticula albifasciella Heinemann, 1871: 222. 2 Syn- types, Germany, West: Braunschweig, e.l. Quer- cus, Heinemann (depository unknown) [not exam- ined]. [Nepticula argyropeza; Stainton, 1854: 300 (partim, imago only); 1859: 433; 1862: 188—191, pl. 9, fig. 2 m (imago); Meyrick, 1895: 726, sbtoniel uon.] Nepticula subapicella Stainton, 1886: 238. Lectotype 3 (here designated), England: Beckenham, Pal- ings, 17.vi.[18]51, S 7609, 57, Stainton, Genitalia slide BMNH 22609 (BMNH) [examined] (Syno- nymised by Emmet, 1974b: 274—276). Nepticula albifasciella; Heinemann & Wocke, 1877: 769; Snellen 1882: 1002; Sorhagen, 1886: 312; Waters, 1928: 248—251 (redescription, biology); Petersen, 1930: 77; fig. 121bis (6 genitalia); Kli- mesch, 1936: 210; Szöcs, 1965: 84. Nepticula subbimaculella var. albifasciella; Rebel, 1901: 228; Meess, 1910: 481. Dechtiria albifasciella; Beirne, 1945: 205, fig. 65 (d genitalia); Emmet, 1971: 246, 247. Stigmella albifasciella; Klimesch, 1951: 66; Gerasi- mov, 1952: 224; Klimesch, 1961: 762; Lhomme, 1963: 1204; Borkowski, 1969: 110. Stigmella (Dechtiria) albifasciella; Hering, 1957: 867 (mine). Trifurcula (Ectoedemia) albifasciella; Johansson, 1971: 245. Ectoedemia (Dechtiria) albifasciella; Borkowski, 1972: fig. 13 (venation). Ectoedemia albifasciella; Bradley et al., 1972: 3; Borkowski, 1975: 491; Emmet, 1976: 199, pl. 6 fig. 10, pl. 12 fig. 30. Trifurcula albifasciella; Karsholt & Nielsen, 1976: 18. [Dechtiria argyropeza; Beirne, 1945: 205, fig. 66 (d genitalia) misidentification.] Diagnosis: only separable from the other members of the complex in the female sex, by the lower number of convolutions in the ductus spermathecae. Externally also very similar to preisseckeri and haraldi, which can however easily be separated on genitalia (see there). Dis- tinguished from E. subbimaculella complex by absence of basal spot, truncate gnathos and sin- gle carinae in male and wider convolutions of ductus spermathecae in female. Other species with white costal and dorsal spot (not metallic) have these spots opposite, or forming an wo straight fascia, and a hair-pencil in male. erythrogenella has a similar pattern, but er: silver spots. Description. Male. Forewing length 2.32—2. 96 mm (2.68 + 0.17, 23), wingspan 5.2—6.4 mm. Head: frontal tuft and collar uniformly orange to fer- ruginous. Antennae with 34—41 segments (36.4 + 1.9, 19). Thorax blackish fuscous, with a few white scales at tip of mesoscutum and tegulae. Forewings blackish fuscous, with a white dorsal spot in middle and a costal spot before middle, sometimes united to form a fascia. Hindwing without hair-pencil, but with costa! bristles. Female (fig. 67). Forewing length 2.32—2.92 mm (2.67 + 0.18, 24), wingspan 5.2—6.5 mm. Antennal segments 25—28 (26.3 + 1.0, 23). Male genitalia (figs. 117, 261, 308, 309, 379). Capsule length 244—321 um (292.1 + 18.5, 13). Tegumen distinctly produced into almost trian- gular, rounded pseuduncus. Gnathos (fig. 308, 309) with central element parallel-sided, with blunt, truncate tip. Valva (fig. 261) length 180— Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 53 236 wm (220.1 + 14.1, 14), apically distinctly narrowed into pointed tip; inner margin strong- ly convex, becoming concave near tip, serrate by prominent sockets of numerous setae on in- ner and dorsal surfaces. Aedeagus (fig. 379) 236—313 um (275.8 + 20.1, 14), carinae point- ed, single. Female genitalia (figs. 191, 192, 416, 451). T7 with a row of 6-12 setae along posterior margin; T8 with two lateral groups of scales and 2—6 setae each; S8 almost quadrate, with parallel sides. Anal papillae with 13—29 setae. Vestibu- lum with vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectina- tions near entrance of ductus spermathecae. Corpus bursae 660—825 um, without pectina- tions; signa dissimilar, longest 460—560 um (519 = 34.7, 10), shortest 395—530 (473 + 39.0, 11), 4.97.5 X as long as wide. Ductus spermathecae with 2/4—2% convolutions, the convolutions being very wide and prominent (fig. 416). Larva. Yellowish white with light brown head-capsule, inconspicuous ganglia. Penulti- mate instars with indistinct brown ventral plates. Biology. Host plants: Quercus robur L., and Q. pe- traea (Mattuschka) Liebl. Occurs on several other deciduous oaks in botanical gardens, and occasionally on Castanea sativa. Mine (fig. 490). Egg on upperside beside vein, or midrib. Mine starting as narrow linear gal- lery, often along midrib and later following lateral vein outwards, abruptly changing into al- most rectangular blotch; sometimes blotch | takes form of wide, irregular gallery. Early mine with linear frass, in blotch frass in basal half. Life history. Univoltine, larvae from end of August until October, usually much earlier than heringi and subbimaculella, but occasionally still feeding in green islands in late October; adults flying in May and June. Distribution (fig. 522). Widely distributed in Central and North Eu- rope, apparently occurring farther northwards than subbimaculella and heringi. In Scandinavia as far north as the limit of Quercus in southern Finland and north of Stockholm in Sweden. Not yet recorded from Norway, but presumably oc- curring along the south coast. Common in Great-Britain as far north as the Scottish High- lands. The distribution in the south is hardly known, due to confusion with other species of the complex. E. albifasciella is there with cer- tainty known from Austria, Hungary and cen- tral Greece (Pindhos mountains). Remarks. This species has been the subject of much confusion. Stainton (1854, 1859, 1862) misiden- tified it as E. argyropeza Zeller, and incorrectly equated the immature stages with those of the real argyropeza. The imago of argyropeza he described as apicella Stainton (see under E. ar- gyropeza). By 1863 Stainton was aware of this incongruency, but did not settle the problem, since he thought that Fritsche was going to pub- lish the solution (Stainton, 1886; Emmet, 1974b). Not until 1886 did he propose the name subapicella for the adults he had previously de- scribed as argyropeza Zeller, still without knowing the life-history. I have studied three specimens from the Stainton collection, labelled as argyropeza which he presumably used for his description of argyropeza and, hence, subapicel- la. From these specimens I selected a lectotype of subapicella. Although Heinemann (1871) also noted Stainton’s misinterpretation of argyropeza, he did not link it up with the new species which he reared from oak, and described as albifasciella. In the Niedersachsisches Landesmuseum Han- nover, there is no material of this species left in the Heinemann collection (pers. comm. R. Jo- hansson), neither in the Berlin or Leningrad museums. However, the clear description, with the note on the foodplant, and the type-locality (Braunschweig) make it most likely that the pre- sent interpretation of albifasciella is correct. Waters (1928) was the first to describe the bi- ology of albifasciella in detail, and to separate it from subbimaculella. Since that time mines and larvae were still often confused with heringi (described in 1934), and in southern Europe with the other species of the complex. I have only seen correct albifasciella females reared from Quercus robur and Q. petraea, all specimens reared from Q. pubescens appear to belong to either E. pubescivora or contorta. However, as this refers to comparatively few specimens, it cannot definitively been concluded that these species are completely host-specific. Material examined: 128 d, 109 2, 23 ex. — Aus- tria: 1 d, Klosterneuburg, Freiberg, 9.v.1932, Preis- 54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 secker; 3 9, Klosterneuburg, Buchberg, el. 8— 16.v.1942, Q. robur, Preissecker (NMW); 1 d, 1 ®, 5 km W. Völkermarkt, Pörtschach (Kärnten), e.l. 27— 30.1v.1984, Quercus robur, J. J. Boomsma & E. J. van Nieukerken (ZMA). — France: 1 6, Pontault, 18.v.1977, Leraut (coll. Leraut). — Germany, West: 1 3, Rohr (Württemberg), el. 18.11.1934, Worz (LNK); 1 6, Schwabisch Hall, 13.vi.1978, W. Speidel (coll. Speidel). — Germany, East: 2 d, Berlin, Fin- kenkrug, 15.v.1923, 22.v.1930, Hering; 1 d, 3 ?, Nordhausen, 24—29.v.1898, Petry (MHUB). — Great Britain: 3 d (lecto- and paralectotypes of suba- picella), Beckenham, Palings, 17 + 22.v1.1851, Stain- ton (BMNH); 1 ©, Saffron Walden, e.l. 25.v.1980, Bryan, Emmet & van Nieukerken; 1 2, Southampton, 15.vi.1935, Fassnidge (ZMA); 2 6, no locality, 8.v.1884, Stevens; 3 6,5 ©, no further data, Walsing- ham (BMNH). — Greece: 1 6, 2 ©, Palaiokastron, Evritania, 1200 m, e.l. 8—13.v.1981, Quercus petraea s.]. 21.1x.1980, Menken & van Nieukerken (ZMA). — Hungary: 1 ©, Budapest, Petnehäzi-rét, el. 20.v.1979, Q. petraea, J. Szöcs; 1 d, 1 2, Matra He- gység, Sástó, el. 12 + 14.v.1973, Q. petraea, Q. ro- bur, J. Szöcs (TMAB). — Netherlands: 98 4,78 2, 23 ex. from following localities: Aerdenhout, Arnhem, Bergen op Zoom, Breda, Bussum, Doetinchem, Drie- sum, Epen, Groesbeek, Den Haag, Helvoirt, Hilver- sum, Hoge Veluwe near Deelen, Hollandse Rading, Horst, Hulshorst, Leeuwarden, Leuvenum, Loenen (Gld.), De Lutte, Nunspeet, Oosterbeek, Overberg, Overveen, Rhenen, Rockanje, Rotterdam, Rijs, Sant- poort, Tietjerk, Ubbergen, Vaals, Venlo, Wageningen, Wassenaar, Winterswijk, Zandvoort (RMNH, ZMA, AFW, coll. Huisman, coll. Kuchlein). — Poland: 3 6, Dabie (Alt Damm), 11.iv.Krone (TMAB); 5 d, 10 2, Krosno Odr. (Crossen a. Oder), e.l. 18.v—10.v1.1930, Quercus robur, Hering; 2 d, Osiecznica (Günters- berg O.), near Krosno, 6.vi.1915, Hering (MHUB). — Switzerland: 1 ®, Lussy (VD), LS 05A, el. 10.vi.1977, S. E. Whitebread (coll. Whitebread). Mines. — On Castanea sativa. — Great Britain: Reading. On Quercus petraea and robur. — Austria: Hof am Leithagebirge; Hundsheimer Berg near Hain- burg; Völkermarkt. — Belgium: Zolder. — France: Andlau. — Germany, West: Blankenheim; Wies- baum. — Great Britain: Little Waltham; Reading; Redhill. — Greece: W. of Palaiokastron, Evritania. — Italy: Tolmezzo. — Netherlands: many localities. Males of albifasciella-complex with uncertain iden- tity. 5 d. — Spain: 1 d, San Ildelfonso (La Granja), 22.vi.1902, Chrétien (MNHN); 1 d, Sierra de Alfa- car, 24.iv.1880, Staudinger (MHUB). — Turkey: 1 6, Asia minor, SW of Yalova, Sea of Marmara, 11.v.1969, Kasy (NMW). — USSR: 1 d, Krasnoar- meysk (Sarepta), 22.v.1859, Christoph (BMNH). — Yugoslavia: 1 4, Drenovo, near Kavadarci (Macedo- nia), 20—30.v.1957, Kasy (NMW). 30. Ectoedemia (Ectoedemia) cerris (Zimmermann, 1944) (figs. 68, 118, 193, 194, 262, 310, 380, 452, 492, 548) Nepticula cerris Zimmermann, 1944: 121. Lectotype ® (here designated), Czechoslovakia: Moravia merid., Lednice (Eisgrub), F. Zimmermann, Geni- talia slide VU 1333 (MHUB) [examined]. Nepticula sp.; Skala, 1942: 6, 7, figs. 1, 2 (description of species, later named montissancti). Nepticula montissancti Skala, 1948: 121, 122. Holo- type, Czechoslovakia: Mikulov (Nikolsburg), v. 1943, e.l., Quercus cerris (Skala) (lost) [not exam- ined]. Syn. nov. Stigmella (Dechtiria) cerris; Hering, 1957: 866, fig. 555 (mine). Nepticula (Dechtiria) cerris; Szöcs, 1968: 227. Ectoedemia cerris; Sz6cs, 1978: 266; 1981: 210. Diagnosis: separated from the other members of the complex by the ductus spermathecae of the female, with 3'!2—4 convolutions. Description. | Male. Forewing length 2.24—2.28 mm (3), | wingspan + 5.0 mm. Antennae with 32-34 (3) segments. Similar to albifasciella, fascia general- ly broken. Female (fig. 68). Forewing length 2.32—2.4 mm (5). Wingspan 5.2—5.3 mm. Antennae with 25—28 segments (4). Male genitalia (figs. 118, 262, 310, 380). As albifasciella. Capsule 250—285 um (2); valva 200—205 um (2); aedeagus 245—250 um (2). Female genitalia (figs. 193, 194, 452). T7 with a row of 6 setae; T8 with 3—6 setae on each side. Anal papillae with 8—13 setae. Corpus bursae 790—860 um; longest signum 462—560 (4), shortest 418—540 (4), 5.4—7 X as wide as long. Ductus spermathecae with 3/2—4 convo- lutions. Larva. Whitish, with dark head-capsule and conspicuous black ventral plates which are shed during final instar. Biology. Hostplant: Quercus cerris L. Mine (fig. 492). Egg on upperside, on or near vein. Early mine narrow gallery, following vein or contorted, with broken linear frass; suddenly widening into large blotch, in which frass 1s ac- cumulated near opening. Mine often away from the midrib. Life history. Univoltine. Larvae have been found from late September to the end of Octo- ber, but most plentiful in early October. The adults appeared in May. Distribution (fig. 548). Known from Hungary, Moravia, eastern Austria, Italy and Yugoslavia. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 55 Remarks. Skala (1948) described montissancti as a third species on Quercus cerris, separate from cerris and liechtensteini, but his description of mine and adult clearly indicate that he was describing cerris again, hence the synonymy. The holotype was according to Skala himself destroyed by psocids. E. cerris is in the autumn the earliest Ectoedemia species mining on Q. cerris. In the first week of October 1983 we found many feeding larvae in Austria, but no other Ectoede- mia species, whereas in the last week of Octo- ber, on the same localities almost only empty mines were found between many larvae of liechtensteini and gilvipennella. Material examined: 7 6, 13 2. — Austria: 5 2, Hof am Leithagebirge, S. of Mannersdorf (Niederöst), e.l. 3—7.v.1984, J. J. Boomsma & E. J. van Nieukerken; 1 ©, Wien, Kahlenberg, SE, 400 m, e.l. 3.v.1984, E. J. van Nieukerken (ZMA). — Czechoslovakia: © lecto- type, see above. — Hungary: 1 4, Budaörs, Csiki-he- gyek, el. 14.v.1971, Q. cerris, J. Szöcs; 2 6,4 ?, Szar, Q. cerris, el. 1.v.1965, 20.1v.1966, 1.v.1966, 18— 19.v.1968, J. Szócs; 1 d, Törökbálint, e.l. 15.v.1965, J. Szócs (TMAB). — Italy: 2 d, P. N. d'Abruzzo, Opi, Bivio, la Camosciara (L’Aquilla), el. 5—7.v.1984, S. B. J. Menken; 1 6, 2 ©, between Tolfa-Allumiere (Roma), e.l. 9—16.v.1984, S. B. J. Menken (ZMA). Mines. — Austria: Hof am Leithagebirge; Eisens- tadt; Loretto, N. of Eisenstadt; Wien, Kahlenberg; — Hungary: Törökbálint. — Italy: Opi; Sabaudia; Tol- fa; Veio. — Yugoslavia: S. of Han Knezica, N. of Pri- jedor. 31. Ectoedemia (Ectoedemia) pubescivora (Weber, 1937) comb. n. (figs. 69, 119, 195, 196, 263, 311, 381, 453, 493, 547) Nepticula pubescivora Weber, 1937b: 212, fig. 2. Lec- totype ® (here designated), Switzerland: Somaz- zo, 12.x.1932, Querc. cerris (sic!), Weber, Genita- lia slide ETH 1236 (ETHZ) [examined]. Stigmella pubescivora; Klimesch, 1948: 73, 74, figs. ; 52—54 (4 genitalia); Klimesch, 1951: 65; Hering, 1957: 870, fig. 547 (mine). Trifurcula (Ectoedemia) pubescivora; Kasy, 1978: 4. Diagnosis: separated from the other members of the complex by the ductus spermathecae in the female, with 6 wide convolutions. Description. Male. Forewing length 2.24—2.56 mm (2.45 + 0,13, 5), wingspan 5.0—5.8 mm. Antennae with 34— 35 (3) segments. Further as albifasciel- la. Female (fig. 69). Forewing length 2.4—2.76 mm (2.55 + 0.11, 9), wingspan 5.2—6.0 mm. Antennae with 25—27 segments (25.7 + 0.8, 7). Male genitalia (figs. 119, 263, 311, 381). As albifasciella. Capsule 270—300 um (3); valva 223—236 um (3); aedeagus 253—274 um (3). Female genitalia (figs. 195, 196, 453). T7 with a row of 6—10 setae; T8 with 2—5 setae on each side. Anal papillae with 10—17 setae. Cor- pus bursae 680—935 um; longest signum 430— 650 um (543 + 46, 14), shortest 395—550 um (485 + 43, 14), 4.3—6 X as long as wide. Duc- tus spermathecae with 5—6 very wide convolu- tions. Larva. As in cerris, with black ventral plates. Biology. Host plant: Quercus pubescens Willd. The specimens in the type-series are labelled Q. cer- ris, but Weber refers clearly to pubescens in his description. Mine (fig. 493). Egg on either surface of leaf. Mine largely as in albifasciella, but both linear part and blotch part often more contorted, and blotch often more forming wide gallery. Life history. Univoltine. Larvae of the type- series have been found in mid October, adults were reared or collected in late May or first half of June. Distribution (fig. 547). With certainty only known from the material examined. The records of mines on Quercus pu- bescens from France and Italy are probably cor- rect. Other records are doubtful, and not in- cluded here. Material examined: 9 6, 20 2. — France: 2 9, “Nesp.” (? near St. Pons, dep. Herault), 15.v1.1904, Chrétien (MNHN); 2 2, Viens (Vaucluse) (near Apt), el. 16—17.v.1979, Quercus pubescens, Buvat (coll. Buvat). — Italy: 3 8, 5 ©, Sardegna, Belvi, environs, 700 m, 29.v—15.v1.1975, F. Hartig (MRST); 4 ©, Sar- degna, Gennargentu, Belvi, 800 m, 19.v.1976, G. Der- ra (coll. Derra): 4 d, 5 2, [Sicilia, Taormina], 572, Groschke (SMNS). — Switzerland: 2 d, 2 © (lecto- and paralectotypes), Somazzo, Monte Generoso, min- es 12.x.1932, Weber (ETHZ). Mines. — France: Aix-en-Provence; Viens (Vau- cluse), leg. Buvat. — Italy: Abruzzi: Alfredena; Goia dei Marsi; Sicilia, Taormina, leg. Groschke (BMNH). — Switzerland: Astano, leg. + coll. Whitebread; So- mazzo, leg. Weber (ETHZ); idem, leg. + coll. White- bread. 32. Ectoedemia (Ectoedemia) contorta sp. n. (figs. 70, 120, 197, 198, 312, 382, 454, 547) Ectoedemia spec.; Van Nieukerken in Kasy, 1983: 5. 56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Ectoedemia ct albifasciella; Van Nieukerken in Kasy, 1983: 5. Type material: Holotype ©, Hungary: Bu- daörs, Csiki-hegyek, Quercus pubescens, e.l. 6.v.1966, J. Szöcs, Genitalia slide VU 1388 (TMAB). Paratypes, 8 9. — Austria: 1 9, Hundsheimer Berg, Porta Hungarica (near Hainburg), 19.vi.1976, F. Kasy; 1 ©, Leithage- birge, N. Burgenland, Zeilerberg S., 30.v.1964, Kasy & Vartan (NMW). — Hungary: 1 9, Csopak, 3.v.1971, Q. pubescens, J. Szöcs; 1 9, Eszakborsodi-karszt, Haragistya, e.l. 3.v.1965, Q. pubescens, J. Szöcs; 1 ®, Matra Hegység, Sástó, el. 16.v.1973, Q. robur, J. Szöcs; 3 9, Nagykovacsi, Kis Szénas (W. of Budapest), e.l. 14—15.v.1964, Q. pubescens, J. Szöcs (TMAB, ZMA). Other material: 4 6, probably belonging to contorta. — Austria: 2 3, Hundsheimer Berg (near Hainburg), 17.vi + 8.vii.1980, F. Kasy (NMW). — Hungary: 1 6, Budaörs. Csiki-he- gyek, el. 10.v.1966, Q. pubescens, J. Szöcs; 1 3, Nagykovacsi, Kis Szenas, e.l. 8.v.1964, Q. pu- bescens, J. Szöcs (TMAB). Diagnosis: easily separated from other fe- males in the species complex by the long spi- raled ductus spermathecae, with 10%—131% convolutions. E. nigroparsella has a similar duc- tus, but has a very different wing pattern. Description. Female (fig. 70). Forewing length: 1.84—2.56 (2.24 + 0.21, 9), wingspan 4.6—5.4 mm. Anten- nae with 22—26 segments (24 + 1.2, 9). Further as albifasciella. Male. Forewing length 2.36—2.48 mm, wing- span 5.2—5.6 mm. Antennae with 32—35 seg- ments. Female genitalia (figs. 197, 198, 454). T7 with a row of 10—12 setae; T8 with 2—5 setae on each side. Anal papillae with 9—21 setae. Cor- pus bursae 715—925 um; long signum 460— 585 um (520 + 52.2, 8), short 430—550 um (487 + 54.1, 8). Ductus spermathecae with 10%—12 (in 1 specimen 1312) convolutions. Further as albifasciella. Male genitalia (figs. 120, 264, 312, 382). Simi- lar to albifasciella. Capsule length 257—278 um. Valva 210—227 um. Aedeagus 257—278 um. Larva not examined. Biology. Hostplants: Quercus pubescens Willd. One specimen reared from Q. robur L. Mine unknown, but since all specimens reared were identified by Szöcs as albifasciella, it probably is very similar to the mine of albifas- ciella. Life history. Univoltine. Adults reared or collected in May, June, and early July. Larvae collected in autumn, but exact data unknown. Distribution (fig. 547). At present only known from eastern Austria and Hungary. Remarks. This species was discovered amongst material identified as albifasciella. All specimens reared by Szöcs from Quercus pubescens appear to be- long to contorta, and all but one reared from Q. robur and Q. petraea are the real albifasciel- la. Only one contorta has been reared from Q. robur. Also the Austrian localities have dense stands of Q. pubescens, so it seems likely that E. contorta is restricted to this oak, and an eastern vicariant of E. pubescivora. As in the other species of this complex, only the females can be identified with certainty, therefore the males are excluded from the type- series, and the order of description is changed accordingly. The Ectoedemia subbimaculella complex The complex of species around E. subbimacu- lella is one of the most difficult species com- plexes in Nepticulidae, and not completely un- derstood. Externally all these species are ex- tremely similar, and show only slight differences in head-colour and size. The male genitalia do not provide constant diagnostic characters and the female genitalia only show minute differences to separate subbimaculella from other species. More than one species has been described because of differences in larval habit and foodplant choice. The larva of E. sub- bimaculella invariably slits its mine open during its last instar, and the larva of E. phyllotomella cuts out a circular disc at the end of its mine. The other species in this complex, without hav- ing such pecularities, have been described be- cause they feed on different species of Quercus, or Castanea, viz. heringi and quercifoliae on Q. robur, and Q. petraea, zimmermanni on Q. pubescens, liechtensteini on Q. cerris and sa- tivella on Castanea sativa. In my experience the larvae found on Q. robur, Q. petraea, Q. pu- bescens and Castanea do not show any differ- ence, but larvae collected on Q. cerris are very EE ES TS us Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 57 different in colour, agreeing with the descrip- tion of liechtensteini. Similar larvae, however, have also been collected in low number on Q. pubescens and Q. petraea, together with the commoner type, so that food plant difference does not seem te be constant. By electrophoresis of allozymes there is indication of some isola- tion in the following species, but in contrast with other situations no diagnostic enzymes have been found: subbimaculella, “heringi” from Q. robur and Q. pubescens and liech- tensteini” from Q.cerris and Q. pubescens (Menken, in preparation). On the ground that the larvae from Castanea and Q. pubescens do not show differences from those from Q. robur, zimmermanni, sativella and quercıfoliae are considered provisionally to be synonymous with beringi. This hypothesis is open to further tests. Hereafter only E. subbimaculella is de- scribed fully, and the other species only in so far as they differ from it. 33. Ectoedemia (Ectoedemia) subbimaculella (Haworth, 1828) (figs. 71, 121, 199, 200, 265, 313, 384, 417, 455, 494, 523) Tinea subbimaculella Haworth, 1828: 583. Lectotype d (here designated), [England], Haworth Coll.; Stainton Coll., Genitalia slide BM 22595 (BMNH) [examined]. Microsetia nigrociliella Stephens, 1829: 208 [nomen nudum]. Microsetia nigrociliella Stephens, 1834: 267. Lecto- type d (here designated), [England], Stephens coll., Genitalia slide BM 22599 (BMNH) [exam- ined]. Syn. nov. Nepticula cursoriella Zeller, 1848: 326. Holotype ?, Germany: Frankfurt am Main, Heyden (deposito- ry unknown) [not examined]. Microsetia subbimaculella; Stephens, 1829: 208; 1834: 267. Nepticula subbimaculella; Stainton, 1849: 29; 1854: 300; 1855: 258—271, pl. 7, fig. 3; Frey, 1856: 379; 1857: 397, 398; Stainton, 1849: 433; Wocke, 1871: 339; 1874: 102; Heinemann & Wocke, 1877: 767; Snellen, 1882: 1002—3; Sorhagen, 1886: 310, 311; Meyrick, 1895: 725, 726; Tutt, 1899: 352; Rebel, 1901: 228; Meess, 1910: 481; Sorhagen, 1922: 56, 57 (partim); Meyrick, 1928: 863; Waters, 1928: 248—251 (differences with albifasciella); Petersen, 1930: 77, fig. 121 (d genitalia); Szöcs, 1965: 86. Stigmella subbimaculella; Klimesch, 1951: 65; Gerasi- mov, 1952: 262; Klimesch, 1961: 761; Lhomme, 1963: 1204; Borkowski, 1969: 111. | Dechtiria subbimaculella; Beirne, 1945: 205, fig. 64 (9 genitalia); Emmet, 1971: 247, 248. Stigmella (Dechtiria) subbimaculella; Hering, 1957: 866, fig. 533 (mine). Trifurcula (Ectoedemia) subbimaculella; Johansson, 1971: 245. Ectoedemia subbimaculella; Bradley et al., 1972: 3; Borkowski, 1975: 490; Emmet, 1976: 200, fig. 60a; b, pl. 7, fıg.,3, pl. 12; fig. 32. Trifurcula subbimaculella; Karsholt & Nielsen, 1976: 18. [no genus] cursoriella; Herrich-Schaffer, [1853]: pl. 106, fig. 844. Nepticula cursoriella; Herrich-Schäffer, 1855: 356. Diagnosis: from most other Ectoedemia spe- cies distinguished by the white basal spot on the forewing and absence of hair-pencil in male. Very difficult to separate from other species in the complex, which have usually a darker head and are slightly smaller. The differences in the male genitalia are not diagnostic. The female can be separated by the wider convolutions in the ductus spermathecae. E. subbimaculella is most easily identified by the dark larval head and pro- thorax and the slit in the mine. Description. Male. Forewing length 2.24—2.8 mm (2.50 + 0.15, 26), wingspan 4.8—6.1 mm. Head: frontal tuft yellowish orange, sometimes with fuscous scales on vertex; collar dark brown. Antennae with 31—36 segments (33.3 + 1.3, 21). Thorax black, with some white scales at tips of meso- scutum and tegulae. Forewing blackish fuscous with a white basal spot along dorsal margin, a dorsal spot in middle and a costal spot before middle, sometimes uniting to form a fascia. Hindwing without hair-pencil, but with costal bristles. Female (fig. 71). Forewing length 2.16—2.8 mm (2.52 + 0.19, 25). Antennae with 24—29 segments (25.7 + 1.1, 24). Male genitalia (figs. 121, 265, 313, 384). Cap- sule length 231—304 um (274.1 + 19.2, 24). Te- gumen produced into rounded pseuduncus. Gnathos (fig. 313) with central element gradual- ly narrowing to rounded tip. Valva (fig. 265) length 193—244 um (222.7 + 13.8, 25), apically gradually narrowed into blunt tip; inner margin little convex to concave, serrate by prominent sockets of many setae on inner and dorsal sur- faces. Aedeagus (fig. 384) 210—261 um (243.5 + 14.3, 23), carinae with variable number of spines. Female genitalia (figs. 199, 200, 417, 455). T7 with a row of 6—10 setae along anterior margin of T8; T8 with two lateral groups of scales and 3—7 setae each; S8 with converging margins. Anal papillae with 9—16 setae. Vestibulum with 58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 vaginal sclerite, a dorsal spiculate pouch, and a group of densely packed pectinations near en- trance of ductus spermathecae. Corpus bursae 450—710 um, without pectinations; signa dissi- milar, longest 390-514 (459.0 + 34.3, 11), shortest 339—467 wm (408.3 + 38.4, 11), 4.4— 5.6 X as long as wide. Ductus spermathecae with 2/4241 (rarely 3) convolutions, wider than in heringi, narrower than in albifasciella (fig. 417). Larva. Translucent glossy white, with dark brown or black head-capsule and prothoracic plate. Ganglia more or less conspicuous. Ventral plates absent. Biology. Host plants: Quercus robur L., Q. petraea (Mattuschka) Liebl., Q. pyrenaica Willd. and Q. pubescens Willd., a few mines known from Oncenriseles in Tagodavt. Rarely on Q. rubra L. In botanical gardens on a wide variety ot de- ciduous oaks. Mine (fig. 494). Egg on upperside of leaf, be- side vein. Mine: narrow linear gallery along vein, abruptly changing in blotch, usually in an- gle between midrib and lateral vein. The larva makes a slit in the under epidermis, through which water and frass fall out of the mine. In Austrian mines on Q. pubescens the slit was of- ten in the upper epidermis or in both surfaces. When the egg is laid along a lateral vein, the lar- va usually feeds towards the midrib. Life history. Univoltine, larvae from late Sep- tember until November, adults flying in June and July Distribution (fig. 523). Widely distributed in West and Central Eu- rope, in Scandinavia only in southern Sweden and Denmark, most northern records being misidentifications (R. Johansson, pers. comm.); it is not recorded from Ireland and Scotland. In the south the distribution is insufficiently known, confirmed records are available from northern Italy, Sicily, Hungary, Yugoslavia and southwest USSR. Remarks. The lectotype is a male in good condition which was placed in the Stainton collection with a “Type” label. On examining the lectotype of nigrociliella Stephens, also from Stainton’s col- lection, the synonymy was confirmed, which was already suggested by several authors (Stain- ton, 1855; Bradley et al., 1972). Types of curso- riella Zeller could not be found, but it is likely to be a synonym of subbimaculella, and has al- ways been treated as such since Herrich- Schaffer (1855). Until the beginning of this century, this was the only oak-mining species of this group rec- ognised by most authors, even albifasciella was generally considered a variety. Waters (1928) was the first to recognise the differences in bi- ology between subbimaculella and albifasciella. Therefore all older literature records are useless, unless a clear description of the characteristic mine with slit is given. More recent records of adults which have not been reared have to be checked since they are easily confused. Hering (1957) mentioned a probable new species from Sicily on Q. pubescens, with similar mines, but with larvae making cocoons in their mines. In BMNH there are such mines, but in all cocoons | | which are still in these mines, pupae of parasitic Hymenoptera can be observed. The phenome- non of parasitised larvae, spinning their cocoons inside the mine has been noted in several spe- cies, thus these are probably subbimaculella mines. This is further corroborated by subbima- culella adults in the Groschke collection, which probably come from Taormina (see also carad- Jai). Material examined: 116 6, 122 ®, 3 ex. — Austria: 11 8, 14 2, Hainburg: Hundsheimer Berg, 200—400 m, el. 8—21.vi.1984, Quercus pubescens, E. J. van Nieukerken (ZMA); 1 dg, Hundsheimer Berg (near Hainburg), 28.vi.1976, F. Kasy; 1 6, Klosterneuburg, Buchberg, el. 14.v.1942, Q.robur, Preissecker (NMW); 6 4,5 2, Loretto, 7 km N. Eisenstadt (Bur- genland), 240 m, e.l. 30.iv—14.v.1984, Quercus pu- bescens, E. J. van Nieukerken; 1 d, Wien, Leopolds- berg, W. of Kahlenberg, 200—400 m, e.l. 2.v.1984, Quercus pubescens, E. J. van Nieukerken (ZMA). — France: 1 d, Pessac-Alouette (Gironde), 3.v1.1934, Le Marchand; 1 d, Mutrécy (Calvados), S. of Caen, 15.vi.1919, Le Marchand (MNHN); 1 2, Mulhouse, Bois de Nonnenbruch, 250 m, 12.v1.1977, S. E. Whitebread (coll. Whitebread); 2 6, 1 2, Ozoir la Ferrière, 30.v.1946, Le Marchand; 1 &, Vaucresson (Hauts de Seine), 17.v1.1946, Le Marchand (MNHN); 1 8, Pontault, 28.1v.1977, P. Leraut (coll. Leraut). — Germany, East: 7 d, 11 ©, Berlin, Finkenkrug, e.l. 25—31.v.1930, Q. robur, Hering (MHUB); 2 9, Nordhausen, 27.v.1898, Krone (TMAB); 6 d, 6 9, Potsdam, e.l. 2—18.v.1900, Hinneberg (MHUB). — Great Britain: 2 6 (lectotypes subbimaculella and nı- grociliella, see above); 3 3, 1 ©, Southampton, 15.vi.1935, Fassnidge; 1 2, Weeley (Essex), Maldon Wood, e.l. 11.vi.1980, Bryan, Emmet & Van Nieuker- ken (ZMA). — Hungary: 1 2, Budapest, Hivos, e.l. 24.v.1956. J. Szöcs (TMAB). — Italy: 3 6, 1 ©, [Sıcı- VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia lia, Taormina], 554, Groschke (SMNS). — Nethet- lands: 65 6, 73 ©, from following localities: Aerden- hout, Amerongen, Arnhem, Bergen (N.H.), Berghem, Breda, Bussum, Doetinchem, Echt, Geulhem, Groes- beek, Den Haag, Helvoirt, Herkenbosch, Hilversum, Hollandse Rading, Horst, Hulshorst, De Lutte, Maarn, Naardermeer, Nunspeet, Olterterp, Ooster- beek, Overberg, Overveen, Rijs, Rotterdam, Sant- poort, Ubbergen, Wageningen, Wassenaar, Winters- wijk, Zandvoort, Zwanewater (RMNH, ZMA, AFW, coll. Huisman, coll. Koster, coll. Kuchlein). — Po- land: 1 6, Dabie (Alt Damm), el. 4.vi. Krone (TMAB). — Portugal: 3 4,3 ©, [San Fiel, Beira Baix- al], 9.v, Quercus toza (= Q. pyrenaica), [Mendes], coll. De Joannis (MNNH). — Yugoslavia: 1 2, Backi Monostor, 4 km S. Bezdan (Vojvodina), el. 5— 7.v.1984, Quercus petraea, J. J. Boomsma & E. J. van Nieukerken; 1 d, Krizisce, 10 km NNW Crikvenica (Hrvatska), e.l. 10.v.1984, Quercus pubescens, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. — On Quercus cerris. — Yugoslavia: NE Bihac. On Quercus petraea. Hungary: Törökbálint. — Yugoslavia: NE Bihac; Baëki Monos- tor, near Bezdan. On Quercus pubescens. — Austria: Gumpoldskirchen; Hundsheimer Berg near Hain- burg; Loretto; Wien, Leopoldsberg. — Italy: Picinis- co; Sicilia, Taormina, leg. Groschke (BMNH).- Yu- goslavia: NNW Crikvenica. On Quercus robur. — Austria: Hof am Leithagebirge. — Belgium: Zolder. — Great Britain: Danbury; Earls Colne; Rainham; Tiptree, Weeley. — Netherlands: many localities. 34. Ectoedemia (Ectoedemia) heringi (Toll, 1934) (figs. 72, 122, 123, 203, 266, 314, 315, 385, 418, 456, 495, 524) Nepticula heringi Toll, 1934a: 1, figs. 3, 4. Lectotype 3 (here designated), Poland: Bydgoszcz, Rynko- wo, e.l. 5.11.1934, Quercus penduculata, Toll, Genitalia slide VU 1408 (IPK) [examined]. | Nepticula quercifoliae Toll, 1934b: 71, 81, pl. 2. Lec- totype 2 (here designated), Poland, Bydgoszcz, Rynkowo, e.l. 18.11.1935, Quercus robur, Toll, Genitalia slide VU 1409 (IPK) [examined] [syno- nymised by Borkowski, 1975]. Nepticula sativella Klimesch, 1936: 208, figs. 10—13. Lectotype ® (here designated), Italy: Teriolis me- rid., Naturno near Merano, e.l. 15—19.v.1935, Castanea sativa, J. Klimesch, Genitalia slide VU 1391 (ZSMK) [examined]. Syn. nov. Nepticula zimmermanni Hering, 1942: 26, fig. Lecto- type 2 (here designated), Czechoslovakia, Libo- chowan (near Litomerice), Elbe, vi.1940, Quercus lanuginosa, F. Zimmermann, Genitalia slide VU 0896 (MHUB) [examined]. Syn. nov. Nepticula heringi; Toll, 1934b: 71; Szöcs, 1965: 86. Stigmella (Dechtiria) heringi; Hering, 1957: 867 (mine). Stigmella heringi; Klimesch, 1961: 761; Borkowski, 1969: 110. 59 Ectoedemia heringi; Borkowski, 1975: 491; Emmet, 1979: 16. Trifurcula (Ectoedemia) heringi; Kasy, 1978: 4; Le- raut, 1980: 49. Nepticula quercifoliae; Klimesch, 1936: 190; Szócs, 1965: 87. Stigmella (Dechtiria) quercifoliae; Hering, 1957: 867 (mine). Stigmella quercifoliae; Borkowski, 1969: 110. Ectoedemia quercifoliae; Bradley et al., 1972: 3; Em- met, 1974a: 108, 147, 148; 1976: 200, fig. 60c, d, pl. 12 fig. 31, pl. 6 fig. 11; Leraut, 1977: 91. Stigmella sativella; Klimesch, 1948: 74—76, fig. 55— 57; Klimesch, 1951: 65. Stigmella (Dechtiria) sativella; Hering, 1957: 256, fig. 165 (mine). Stigmella zimmermanni; Klimesch, 1951: 65; 1961: 761. Stigmella (Dechtiria) zimmermanni; Hering, 1957: 866, fig. 540 (mine). Nepticula zimmermanni; Szöcs, 1965: 86. Trifurcula (Ectoedemia) zimmermanni; Kasy, 1978: Ato Ectoedemia zimmermanni; Szöcs, 1981: 210. Klimesch, 1961: 761; Diagnosis: distinguished from E. subbimacu- lella by the darker head and the ductus sperma- thecae in the female; the species is slightly smaller than subbimaculella. Adults not separa- ble from phyllotomella or liechtensteini. In the mine there is no slit, which makes it very similar to the mine of E. albifasciella, however, heringi usually feeds towards the midrib. Description. Male (fig. 72). Forewing length 1.88—2.4 mm (2.18 + 0.18, 14), wingspan 4.2—5.3 mm. Head: frontal tuft ferruginous, on vertex brown to black, a sharp delimitation of the light and dark area at the level of antennal insertion; col- lar similar to vertex. Antennae with 29—32 (—36) segments (31 + 2.0, 13). Thorax and fore- wing as in E. subbimaculella, but basal spot of- ten larger. Hindwing with costal bristles. Female. Forewing length 1.88—2.44 mm (2.14 + 0,18, 8). Antennae with 22—25 seg- ments 23.4 =-71377): Male genitalia (figs. 121, 123, 266, 314, 315, 385). Capsule 230—270 um (249,6 + 14.6, 12). Tegumen broadly rounded, slightly less produc- ing than in subbimaculella. Gnathos (figs. 314, 315) with rather short and broad, rounded cen- tral element. Valva (fig. 266) length 175—215 um (195.4 + 10.2, 12), tip blunt, broader than in subbimaculella, inner margin straight, or hardly convex in proximal third, concave apically. Ae- 60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 deagus (fig. 385) 205—255 um (228.6 + 14.3, 12). Several specimens are not separable from subbimaculella. Female genitalia (figs. 203, 418, 456). T7 with a row of 6—10 setae along posterior margin. T8 with two lateral groups of scales and 1—3 setae. Anal papillae with 9—15 setae. Corpus bursae 410—660 um; longest signum 347—463 um (395.5 + 35.9, 11), shortest 309—420 um (350.1 + 33.6, 10), 4.45.8 X as long as wide. One specimen with much smaller signa: 257, 287 um. Ductus spermathecae with 2—2V4 narrow con- volutions (fig. 418). Larva. Translucent yellowish white, or green- ish white, with dark brown head-capsule. Gan- glia usually conspicuous, but sometimes less so. Ventral plates absent. Separated from albifas- ciella by darker head. Biology. Host plants: Quercus robur L., Q. petraea (Mattuschka) Liebl., Q. pubescens Willd., Q. faginea Lam. and Castanea sativa Miller. Mine (fig. 495). Egg on the upperside beside a vein, often the midrib. Mine starts as narrow linear gallery following vein, usually towards midrib, abruptly changing into a blotch, or false blotch, without slit, usually in angle between midrib and lateral vein. Sometimes the last part resembles more a wide gallery than a blotch. Life history. Univoltine, larvae from late Sep- tember until November, but in southern Spain also found in February, adults flying in May in the south and in June and July more in the north. Distribution (fig. 524). Due to confusion with subbimaculella and al- bifasciella insufficiently known. Apparently lacking in Scandinavia and the Netherlands, scarce in south east England, more common in central Europe. Remarks. This species seems to have the widest range of foodplant species within the subbimaculella group. Some of the synonyms listed here were described as separate species only on the basis of a different foodplant species. These forms, E. zimmermanni on Q. pubescens and E. sati- vella on Castanea sativa, of which lectotypes have been selected, differ neither morphologi- cally, nor biologically and can therefore only be treated as one species. E. heringi and quercifo- liae were both described in 1934, but which was published first is not clear, however, most likely heringi should take priority, since it is also men- tioned in Toll (1934b), as an established species. In this paper Toll compares the larval characters and the mines of both species. N. quercifoliae was originally only described from mines and larvae which were collected in the autumn of 1934. From these, in fact the syntypes, he reared adults in 1935, which can therefore be regarded as type material. The & in Toll’s collection, bearing the label “type”, is selected as lectotype. Material examined: 72 d, 76 © : reared from Quer- cus robur or petraea: 33 6, 25 2. — Austria: 2 6, Klosterneuburg, Freiberg, el. 9.v.1932, 18.1v.1938, Preissecker; 1 ©, Klosterneuburg, Buchberg, el. 25.v.1941, Preissecker (NMW). — France: 3 6, 1 9: Andlau (Bas-Rhin), Kastelberg, el. 9—19.v1.1979, Q. petraea, E. J. van Nieukerken (ZMA). — Hunga- 1 ©, Szentpéterfölde, el. 25.v.1969, Q. robur, J. . Szócs (TMAB). — Poland: 3 d (lecto- and paralecto- types of heringi), Bydgoszcz, Rynkowo, el. 28.1— 7.11.1934, Q. pedunculata, Toll (ZMC, MHUB); 2 d, 1 2 (lecto- and paralectotypes of quercifoliae), same locality, el. 16—18.111.1935, Q. robur, petraea, Toll PAR) DIS, Zi Aelen El wile, O. pezzo, Toll (PAK, MHUB, MNHN). — Yugoslavia: 2 d, S. of Han Knezica, 11 km N. of Prijedor (Bosna), e.l. 25.iv—1.v.1984, Quercus robur, J. J. Boomsma & E. J. van Nieukerken (ZMA). Reared from Q. pubescens: 23 3, 35 ©. — Austria: 3 6,7 2, Hainburg, Hundsheimer Berg, 200—400 m, el. 27.iv.—1.v.1984, E. J. van Nieukerken; 3 6, 8 9, Wien, Leopoldsberg, W. of Kahlenberg, 200—400 m, el. 6—12.vi.1984 (ZMA). — Czechoslovakia: 14 dg, 16 2 (lecto- and paralectotypes of zimmermannı), Li- bochowan (near Litomerice), Elbe, e.l. vi.1940, Zim- mermann (MHUB, ZMC). — Hungary: 1 6, 1 2, Pécs Mecsek, Misina, e.l. 27—29.1v.1966, J. Szöcs; 2 4, 3 2, Törökbälint (W. of Budapest), el. 12— 17.v.1974, J. Szöcs (TMAB). Reared from Quercus faginea: 1 ?, Spain: 3 km NW. San Pedro de Alcantara (Malaga), 300 m, mine 6.11.1984, el. 25—26.iv.1984, E. J. van Nieukerken (ZMA). Reared from Castanea sativa: 2 6,2 2. — Italy: 2 3,1 2 (lecto- and paralectotypes of sativella), Natur- no, near Merano, e.l. 15—24.v.1935, Klimesch (ZSMK); 1 ®, Trento, e.l. v.1946, J. Klimesch (MNHN). Reared from unknown Quercus or not reared, but likely to be heringi: 14 d, 13 9. — Austria: 3 d, 6 &, Hackelsberg, N. of Neusiedlersee, 1971—1977, F. Kasy; 1 ©, Hundsheimer Berg (near Hainburg), 28.vi.1976, F. Kasy (NMW). — France: 2 6, no data, De Joannis (MNHN). — Germany, West; 1 d, 1 9, Stuttgart, Lindental, e.l. 27.iv—4.v.1947, Worz; 1 6, 1 2, Stuttgart, Wildpark, e.l. 9.v.1938, Wörz (LNK); 1 2, Wolfenbuttel, [Heinemann], coll. Staudinger (MHUB). — Germany, East: 1 6, Altenburg, 1874, Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 61 Krause; 1 d, Dresden, Staudinger (MHUB). — Hun- gary: 1 6, Budapest, Zanoshegg, el. 15.v.1960, J. Szöcs; 1 d, Szigetszentmiklos, el. 23.v.1955, J. Szöcs (TMAB). — Poland: 3 4,3 ©, Wroclaw (Breslau), e.l. iv.1869, Wocke (MHUB). Identity uncertain: 1 d, 1 2. — Albania: 1 d, Kula Ljums, 7—14.vi.1918, Alban. Exped. — Yugoslavia: 1 2, Drenovo near Kavadarci, 20—30.v.1957, Kasy (NMW). Mines. — On Quercus faginea. — Spain: Istan; NW of San Pedro de Alcantara. On Quercus petraea. — France: Andlau. — Hungary: Törökbálint. — Po- land: Bydgoszcz, leg. Toll (BMNH). — Yugoslavia: 11 km NE Bihac; Slavonska Pozega. On Quercus pu- bescens. — Austria: Gumpoldskirchen; Hundsheimer Berg; Loretto; Wien, Leopoldsberg. — Czechoslova- kia: Libochowan, near Litomerice, leg. Zimmermann (BMNH). — Hungary: Budaörs. On Quercus robur. — Great Britain: S. of Weeley. — Poland: Bydgoszcz, leg. Toll (BMNH). — Yugoslavia: Han Knezica, N. of Prijedor. 35. Ectoedemia (Ectoedemia) liechtensteini (Zimmermann, 1944) (figs. 124, 204, 496, 525) Nepticula liechtensteini Zimmermann, 1944: 119— 121, fig. 8. Lectotype © (here designated), Czechoslovakia: Moravia merid., Lednice (Eis- grub), F. Zimmermann, Genitalia slide 4775 (MHUB) [examined]. Stigmella (Dechtiria) liechtensteini; Hering, 1957: 866, fig. 558 (mine). Ectoedemia liechtensteini; Szöcs, 1978: 266. Diagnosis: adults cannot be separated from heringi. Larvae intensely amber-yellow, with- out visible ganglia in contrast with greenish white larvae of heringi, which usually have dis- tinct ganglia. Specific status doubtful. Description. Male. Forewing length 2.12—2.16 (3), wing- span 4.8 mm. Antennae with 28—31 segments. Further as heringi. Female. Forewing length 1.8—2.28 (3), wing- span 4.4—5.2 mm. Antennae with 22—24 seg- ments. Male genitalia (fig. 124). Similar to heringi. Capsule length 249 um (2). Valva length 180— | 210 um (2). Aedeagus 223—231 um (3). Female genitalia (fig. 204). T7 with a row of | 6—8 setae. T8 with 3—5 setae at each side. Anal papillae with 10—12 setae. Corpus bursae 460—595 um; longest signum 334—411 um (2); | shortest 291—356 um (2), 4.5—4.9 X as long as wide. Ductus spermathecae with 2—2 Y, incon- spicuous convolutions. Larva. Intensely, glossy amber yellow, with very light brown head-capsule and prothoracic plate. Not the slightest indication of ganglia. Ventral plates absent. Biology. Hostplants. Quercus cerris L. on which it can be very abundant. Very occasionally on Q. pe- traea (Mattuschka) Liebl. or GQ. pubescens Willd. (see remarks). Mine (fig. 496). Egg on leaf upperside. Mine completely similar to heringi, in the axil of the midrib and a lateral vein. Life history. Univoltine. Larvae in October- November, usually much later than E. cerris, es- pecially abundant in late October. Adults (reared) from April to June. Distribution (fig. 525). With certainty from Moravia, east Austria, Hungary and Yugoslavia. Remarks. The separate identity of this species is uncer- tain. Adults are similar to heringi, but the larvae are very different, and can easily be distin- guished. Moreover, larvae of liechtensteini are usually found on Q. cerris, whereas sympatric heringi occurs on other oak species, but never on cerris. However, in autumn 1983 I also found one larva of the liechtensteini type on Q. petraea, in a locality with numerous liechtensteini on Q. cerris, and several larvae on Q. pubescens in Gumpoldskirchen. In the latter locality no Q. cerris grew, but on the Q. pubes- cens some “normal” heringi larvae were also noted. S. Menken (pers. comm.) could find no difference in their allozymes and allozyme dif- ferences with heringi were insignificant. It will be necessary to set up foodplant choice and hy- bridisation experiments in order to solve prob- lems of isolation in this species complex. The striking differences in the larva lead me to consider liechtensteini tentatively as a sepa- rate taxon, having no evidence to the contrary. Material examined, 22 6, 22 2. — Austria: 7 6, 9 2, Hof am Leithagebirge, S. of Mannersdorf (Nie- deröst.), 200 m, el. 2.v, 10—18.vi.1984, Quercus cer- ris, E. J. van Nieukerken; 3 d, 3 2, Loretto, 7 km N. Eisenstadt (Burgenland), 240 m, el. 5—25.v.1984, Quercus cerris, E. J. van Nieukerken; 1 d, Wien, Kahlenberg SE., 400 m, e.l. 30.iv—1.v.1984, Quercus cerris, E. J. van Nieukerken (ZMA). — Czechoslova- kia, 5 4,3 9 (lecto- and paralectotypes), Moravia me- rid., Lednice (Eisgrub), Zimmermann (MHUB, ZMC). — Hungary: 2 4,2 2, Törökbálint (W. of Bu- 62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 dapest), e.l. 5, 11.v.1968, 14, 18.v.1974, Q. cerris, ]. Szócs (TMAB); 2 d, Törökbálint, Nagy-erdö, 5 km N. Erd, el. 25.iv—1.v.1984, Quercus cerris, J. J. Boomsma & E. J. van Nieukerken (ZMA). — Yugo- slavia: 2 6, 4 ©, Backi Monostor, 4 km S. Bezdan (Vojvodina), e.l. 25.iv—4.v.1984, Quercus cerris, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. On Quercus cerris. — Austria: Eisenstadt; Hof am Leithagebirge; Loretto. — Czechoslovakia: Lednice (Eisgrub), leg. Zimmerman (BMNH). — Hungary: Törökbálint. — Yugoslavia: Han Knezica, N. of Prijedor; Baëki Monostor, S. of Bezdan. On Quercus petraea. — Hungary: Törökbalınt (1 mine). On Quercus pubescens. — Austria: Gumpoldskir- chen. 36. Ectoedemia (Ectoedemia) phyllotomella (Klimesch, 1946) comb. n. (figsa/35 1255 20552675 3865497549752) Stigmella phyllotomella Klimesch, 1946: 166, fig. 7, pl. 12. Lectotype d (here designated), Italy: Ligu- ria, Altare near Ferrania, el. 26.1v—7.v.1945, Quercus cerris, 2.x1.1944, Zucht 507, J. Klimesch, Genitalia slide Kl. 270 (ZSMK) [examined]. Stigmella phyllotomella; Hering, 1957: 855 (mine). Diagnosis: adults not separable from heringi, ‘although head slightly lighter. Female separated from subbimaculella by narrower convolutions of ductus spermathecae. Mines very characteris- tic by circular “cut-out”. Description. Male (fig. 73). Forewing length 2.16—2.24 mm, wingspan 4.9—5.2 mm. Antennae with 30—34 segments. Head: frontal tuft yellowish orange, on vertex fuscous. Further as subbima- culella. Female. Forewing length 2.04 mm, wingspan 4.6 mm. Antennae with 23 segments. Male genitalia (figs. 125, 267, 386). Similar to subbimaculella. Capsule length 233—253 um (3). Valva (fig. 267) length 193—210 um (3). Aedeagus (fig. 386) 214—236 um (2). Female genitalia (figs. 205, 457). T7 with a row of 8 setae. T8 with 2—5 setae at each side. Anal papillae with 8—9 setae. Corpus bursae 515—530 um; longest signum 386—390 um, shortest 339—356 um, 4.6—4.9 X as long as wide. Ductus spermathecae with 2 very incon- spicuous convolutions. Larva not examined. Biology. Hostplant: Quercus cerris L. Mine (fig. 497). Egg on leaf upperside, against midrib. Early gallery narrow, following vein or midrib; later becoming highly contorted gallery with linear frass, often forming false blotch. The larva cuts out an oval case from the end of the mine, in which it pupates. The case does not fall immediately to the ground, but after some time, by weathering of the leaf. Life history. Univoltine. Larvae collected in late October and early November, adults reared in April and May. Distribution (fig. 525). Only known from Italy: Liguria and Lucania. Remarks. The peculiar habit of the larva, and the food- plant, suggest that phyllotomella is a separate entity, isolated from the other species of the complex. Study of larvae and electrophoresis of allozymes might shed some light on the degree of genetic isolation from its relatives. Material examined: 3 6, 2 9. — Italy: 2 4,1 9 (lecto- and paralectotypes), Liguria, Altare near Fer- rania, el. 26.iv—7.v.1945, J. Klimesch (ZSMK); 1 6, 1 2, Lucania, Mte Vulture, Laghi di Monticchio, 750 m, e.l. 2—7.1v.1966, F. Hartig (LNK). Mines. — Italy: Ferrania, Ligur. Appenin, leg. Kli- mesch (BMNH) (2 mines only). 37. Ectoedemia (Ectoedemia) spec. (specimen 1375) (figs. 74, 206, 458) Material: 1 2, Iran: 100 km W. Shiraz, 18.1v.1970, Exp. Mus. Vind., Genitalia slide VU 1375 (NMW). Undoubtedly a new species, which I do not name here, because of limited material and lack of knowledge on biology. It is externally most similar to gilvipennella. Description. Male unknown. Female (fig. 74). Forewing length 2.4 mm, wingspan 5.3 mm. Head: frontal tuft ochreous- white; collar white. Antenna with 23 segments. Thorax and forewings uniform light brown irrorate with yellowish white. Female genitalia (figs. 206, 458). T7 with a distinct row of 14 setae along posterior margin. T8 with two groups of few (3—5) setae, no scales. Anal papillae with 17—18 setae. Vestibu- lum with vaginal sclerite, a spiculate pouch with many spines and a dense patch of pectinations near entrance of ductus spermathecae. Corpus bursae 595 um, without pectinations; signa dis- similar, longest 437 um, shortest 360 um, 3.9 x as long as wide. Ductus spermathecae with 3 narrow convolutions. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 63 The Ectoedemia terebinthivora group 38. Ectoedemia (Ectoedemia) terebinthivora (Klimesch, 1975) comb. n. (figs. 75, 126, 201, 202, 268, 316, 383, 412, 459, 498, 540) Trifurcula (Ectoedemia) terebinthwora 1975b: 19—23, figs. 27—33. Syntypes, 4 d, 8 9, Anatolia: Kanlidivane, along road Silifke-Mersin, larvae 31.v.1970, e.l. 24—30. vi.1970, Klimesch (ZSMK) [not examined]. Trifurcula (Ectoedemia) terebinthivora; Klimesch, 1978: 251, figs. 26—28 (mine, d, © genitalia). Diagnosis: externally characterised by small size, light brown ground-colour with yellowish tinge and in male by hindwing almost complete- ly covered with brown androconial scales. E. aegilopidella has similar scales in male but has also a hair-pencil which is absent in terebin- thivora. Description. Male. Forewing length 1.88—2.24 mm (2.08 + 0.12, 8), wingspan 4.1—5.0 mm. Head: fron- tal tuft very variable, from completely yellowish to dark brown, variation not sex-linked; collar similar or slightly lighter. Antennae with 39— 41 segments (40.3 + 1,0, 7). Thorax and fore- wings brown, with an obvious yellow tinge; thorax sometimes apically lighter; forewing with a medial yellowish fascia, somewhat irreg- ular, outer margin concave, sometimes fascia in- distinct. Hindwing covered in basal two thirds with brown lamellar androconial scales, not ex- tending in fringe; costal bristles or hair-pencil absent. Underside forewing with few similar brown scales near base. Female (fig. 75). Forewing length 2.12—2.32 mm (2.23 + 0.09, 8), wingspan 4.7—5.2 mm. Antennae with 33—35 segments (34.1 + 0.7, 7). Male genitalia (figs. 126, 268, 316, 383, 412). Capsule length 197—214 um (4). Tegumen pro- duced into broad, truncate pseuduncus (fig. 412). Gnathos (fig. 316) with very short, round- ed central element. Valva (fig. 268) length 146— 163 um (4), inner margin almost straight, except basally, tip pointed. Aedeagus (fig. 383) 279— 300 um (4), much longer than capsule, with pair of single, pointed, dorsal carinae. Female genitalia (figs. 201, 202, 459). T7 without row of setae. T8 with two lateral patches of scales and setae (6—7). Anal papillae with 8—10 setae. Anterior apophyses remark- ably widened in middle. Vestibulum with vagi- nal sclerite and dorsal spiculate pouch with Klimesch, - many pointed spines, and a dense patch of pecti- nations near entrance of ductus spermathecae. Corpus bursae 470—530 um, covered with minute pectinations, except anteriormost part; signa dissimilar, longest 369—403 um (4), shortest 309—334 um (4), 4.2—5.0 X as long as wide. Ductus spermathecae with 2—2 Y, convo- lutions. Larva. Yellowish white to whitish, in mine appearing greenish, first 4 ganglia distinct. Head-capsule brown. Penultimate stages with 12 ventral brown plates. Biology. Hostplant. Pistacia terebinthus L. Mine (fig. 498). Egg always deposited on leaf underside, close to midrib or lateral vein. Early mine much contorted with thin brownish linear or dispersed frass; later widening into large ir- regular, elongate blotch with dispersed brown frass. Life history. Probably bivoltine, or at least partly. Larvae in late May and June (Klimesch, 1975b) and in September. Adults reared in June and July (from May and June larvae) and May— June (from September larvae). Therefore Kli- mesch’s assumption that the species is univol- tine seems to be incorrect. Distribution (fig. 540). Greece, Ionian and Aegean Islands and Anat- olia. Probably widespread in eastern Mediterra- nean. Record from Keffalinia from mines in old herbarium specimen of Pistacia in Rijksherbari- um, Leiden, no. 897, 363—722. Material examined: 12 d, 12 2. — Greece: 1 9, Athina (Atena), 16.vi.1980, Leo Kohonen (ZMUO); 11 6,9 ©, 3 km E. of Dhelfoi (Fokis), 700 m, el. 2.v—11.vi1.1981, Pistacia terebinthus, 27.1x.1980, S. B. J. Menken, E. J. van Nieukerken (ZMA, BMNH, ZSMK); 1 6, 1 2, Kardhamili (Messinia), a.s.l., el. 14—16.vii.1984, E. J. van Nieukerken (ZMA). — Turkey: 1 ®, Asia minor, Tekir Tepisi, Taurus, 13.vii.1965, Arenberger (LNK). Mines. — Greece: Parnis Oros (Attika); Evvoia: SE Gouvés; Oiti Oros, SW Ipáu (Fthidtis); Dhelfoi (Fökis); Kardhamili (Messinia). The Ectoedemia angulifasciella group This is a rather heterogenous assemblage of Rosaceae mining species, comprising a tight group — hexapetalae, angulifasciella complex, mahalebella and spinosella — and some aber- rant species which at present cannot be included in any other group. 64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 The adults usually have a shining metallic fas- cia, and males have a hair-pencil, or this is sec- ondarily lost. Except in the first three species, the gnathos is divided, and the basal part has a serrate margin. The aedeagus has one pair of carinae, often with additional spines. The valva is comparatively uniform, with a more or less straight inner mar- gin. In female genitalia the vaginal sclerite is pre- sent in most species except spiraeae and agrimo- niae, but the spiculate pouch is less distinct than in previous groups or even absent. The bursa is covered with pectinations. The larvae make gallery-blotch mines, and only the species in the angulifasciella complex have ventral plates in the penultimate stages. Species belonging to this group occur also in Japan and probably in North America (E. rubi- foliella (Clemens)). 39. Ectoedemia (Ectoedemia) erythrogenella (de Joannis, 1908) (figs. 76, 128, 129, 207, 269, 317, 387, 460, 499, 528) Nepticula erythrogenella J. de Joannis, 1908a: 327, 328. Lectotype d (here designated), France: Van- nes, L. de Joannis, Genitalia slide VU 946 (MNHN) [examined]. [Nepticula rubivora; Walsingham, 1891: dentification] Nepticula erythrogenella; J. de Joannis, 1908b: 823, figs. 1, 2, pl. 15 fig. 12 (mine, adult, larva); Kli- mesch, 1940b: 190. Stigmella erythrogenella; Gerasimov, 1952: 238; Her- ing, 1957: 908 (mine); Lhomme, 1963: 1192. 152, misi- Ectoedemia (Dechtiria) erythrogenella; Emmet, 1974c: 129, 130, fig. (mine). Ectoedemia erythrogenella; Emmet, 1976: 195, fig. 59, pl. 9 fig. 16. Trifurcula (Dechtiria) erythrogenella; Gustafsson, 1981b: 466—468, fig. 8 (d, @ genitalia, larva, mine). Stigmella erythrogenella ab. juncta Dufrane, 1949: 9. Diagnosis: separated from all other Rosaceae feeding Ectoedemia by costal spot (or costal part of fascıa) placed distinctly before middle of forewing; in addition separated from angulıfas- ciella complex by absence of hair-pencil in male. Externally similar to albifasciella-complex and preisseckeri, but separated by shining silver spots on forewing and absence of costal bristles in male. Male genitalia characterised by shape of valva, with almost posteriorly directed tip, and undivided, smooth gnathos. Description. Male (fig. 76). Forewing length 1.76—2.28 mm (2.05 + 0.19, 13), wingspan 4.1—5.0 mm. Head: frontal tuft ferruginous, or orange, sometimes becoming fuscous towards crown; collar yellowish white, lighter than frontal tuft. Antenna with 33—41 segments (36.1 + 2.3, 10). Thorax and forewings blackish, with shining sil- very white spots, one slightly before middle on costa, one in middle on dorsum, with sometimes a small spot in between, less commonly united to form a fascia (ab. juncta). Hindwing without hair-pencil or costal bristles. Female. Forewing length 1.88—2.52 mm (2.23 + 0.22, 12), wingspan 4.1—5.6 mm. An- tennae with 25—30 segments (27.5 + 1.4, 8). Male genitalia (figs. 128, 129, 269, 317, 387). Capsule length 189—223 um (206.6 + 13.7, 5). Tegumen distinctly produced into slightly trun- cate pseuduncus. Gnathos (fig. 317) with broad- ly spatulate, undivided, smooth central element. Valva (fig. 269) length 150—180 um (158.6 + 13.2, 5), gradually narrowing into pointed tip, which points almost posteriorly; inner margin approximately straight. Aedeagus (fig. 387) 223—253 um (238.2 + 14.1, 5), with pointed, single carinae. Female genitalia (figs. 207, 460). T7 with a distinct row of 4—10 long setae along posterior margin. T8 trapezoid, with two lateral patches of scales and 3—5 setae. Anal papillae with 6— 11 setae. Vestibulum with vaginal sclerite, a spi- culate pouch (sometimes indistinct) and a dense patch of pectinations near entrance of ductus spermathecae. Corpus bursae 440—690 um, covered with pectinations, except anterior part, especially closely set near vestibulum; signa similar, 300—369 um (326.8 + 24.9, 12), 3.9— 5.6 X as long as wide. Ductus spermathecae with 2 4—3 convolutions. Larva. Dirty grey, but more yellowish in ear- ly stages; ganglia conspicuous. Head capsule dark brown. Ventral plates absent. Biology. Hostplant. Rubus fruticosus L. sensu lato, es- pecially on evergreen Rubus ulmifolius Schott. Mine (fig. 499). Egg on upperside against midrib or vein. Early mine narrow gallery, fol- lowing vein, often turning back, completely filled with blackish frass; finally widening into elongate blotch, with dispersed black frass in basal part, or at sides. Leaves often stained red around mine. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia Life history. Univoltine, with a very long pe- riod of larval feeding. In the northern part of its range larvae from September until November, but in the south larvae can be found all over the winter until March, April and occasionally later. Some data: mid-October, Trieste, many early instar larvae, full-fed after two or three weeks; early February, south Spain, many early instar, fewer late instar larvae, completing their larval cycle in two to four weeks; late March, Sicily, few larvae left; late April, Aures mountains in Algeria, few larvae left, but still giving rise to adults; July, southern France, very few larvae, no adults reared. It is not clear if the July larvae belonged to the old generation or were just very early larvae of the new generation, but since no young larvae were present it is most likely that they belonged to the past generation and were late because of parasitism. Adults emerged in May—July, whether from autumn or early spring larvae. Distribution (fig. 528). Essentially a mediterranean species, which is abundant and widely distributed throughout the ‚ mediterranean region, both along coast and in- land, although it has still to be recorded from many places. Distributed along French Atlantic coast as far as the south coast of England, where it can only be found within a short distance of the sea (Emmet, 1976), as a consequence of its supposed vulnerability to frost. The species has been recorded from Switzerland, where it might occur in Tessin, but it certainly does not occur in Austria as erroneously indicated by Emmet (1976) (Klimesch, in litt.). Material examined: 21 d, 23 9. — Algeria: 1 9, Aures, Dj. Chelia, northern slopes, 1500 m, e.l. 5.vi.1980, Rubus ulmifolius, 29.iv, E. van Nieukerken, G. Bryan, P. Oosterbroek (ZMA). — Cyprus: 3 d, 3 2, Limassol, Yermassoyia, 24 + 28.11.1980, Rubus, B. Gustafsson (RMS). — France: 2 6, 4 ©, Cannes, el. 27.v—12.vi.1889, Rubus fruticosus, ili. Walsing- ham (BMNH); 5 d, 6 @ (lecto- and paralectotypes), Vannes, ronce, 24.vi, 1.vii, Joannis (MNHN, MHUB); 1 6, 3 2, Vannes, el. 28.vi—29.v11.1910, mine 8.x.1909, Joannis, coll. Dufrane (IRSN). — Great Britain: 3 d, 1 ©, Portland, Church Ope Cave, el. 10—17.vi.1982, Rubus fruticosus, 28.ix.1981, Bry- an & Menken (ZMA). — Italy: 1 6, Sicilia (Caltani- setta), W. of Manzarino, e.l. 2—4.v.1981, Rubus ulmi- folius, 25.11.1981, E. J. van Nieukerken (ZMA). — Spain: 1 d,2 ®, 7 km NW San Pedro de Alcantara (Malaga), 350 m, e.l. 21.iv, 15.v, 3—4.vii.1984, Rubus ulmifolius, E. J. van Nieukerken; 4 &, Sierra Blanca, 6 km N. Marbella (Malaga), El Mirador, 800 m, e.l. 65 12.1v—18.v1.1984, Rubus ulmifolius, E. J. van Nieu- kerken (ZMA). — Yugoslavia: 1 d, 3 ®, 7 km SE Piran, Cedle (Slovenia), 300 m, e.l. 22.iv—14.v.1984, Rubus ulmifolius, J. J. Boomsma, E. J. van Nieuker- ken (ZMA). Mines. — Algeria: Aurés, Arris, 32 km SSE Batna; Aurès, Dj. Chélia; La Calle (El Kala); E. of Morris. — Corsica: Pisciatella; Porticcio (near Ajaccio). — Cy- prus: Limassol, Yermassoyia (RMS). — France: Ba- nyuls; Port Vendres; Douelle (Lot), Le Carriol (BMNH); Bretagne (Cotes du Nord) (BMNH). — Great Britain: Harwich; Newhaven (Sussex), Emmet; Portland; St. Osyth. — Greece: Kardamyli (Messi- nia). — Italy: Frascati (BMNH); Roma Fiumicino; Sasso di Bordighera (BMNH); Trieste; Sicilia, Mazza- rino; Sicilia, Montallegro; Sicilia, Taormina (BMNH). — Spain: Marbella; San Pedro de Alcantara; Tunisia: Ain Draham; Hammam Lif; Tabarka. — Yugoslavia: Piran; Rovinj (BMNH). 40. Ectoedemia (Ectoedemia) spiraeae Gregor & Povolny, 1983 (figs. 77, 127, 204, 271, 318, 388, 416, 500, 549) Ectoedemia spiraeae Gregor & Povolny, 1983: 174— 177, figs. 4—7, 9. Holotype d, Czechoslovakia: Ciganka Hill near Murän, 930 m, 26.1x.1981, el. 11.1982, Spiraea media, Gregor & Povolny (De- partment of Entomology, Moravian Museum, Brno) [not examined]. Stigmella sp.; Povolny & Gregor, 1952: 237, figs. c, d (mine). Stigmella spireae (sic!) Gregor & Povolny, 1955: 124, 127 (nomen nudum, no description); Hering, 1957: 1021 (mine). Nepticula spireae; Szöcs, 1968: 229. Diagnosis: externally characterised by light head and collar, almost straight non-metallic fascia and in male yellowish-white hair-pencil and white tuft on underside forewing. Male genitalia characterised by aedeagus without ca- rinae and valvae with serrate inner margin and inconspicuous tip. Female genitalia character- ised by absence of both vaginal sclerite and spi- culate pouch, and by dissimilar signa. Description. Male. Forewing length 2.42—2.52 mm (4), wingspan 5.0—5.6 mm. Head: frontal tuft and collar yellowish-orange. Antennae with 34—36 segments (4). Thorax and forewings blackish, with medial, almost straight, non-shining fascia, often interrupted. Underside of forewing with a tuft of white hair-scales arising near costal reti- naculum and a large scaleless area. Hindwing with a yellowish-white hair-pencil. Female (fig. 77). Forewing length 2.2—2.32 m (2.25 + 0.04, 7), wingspan 4.8—5.4 mm. 66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Antennae with 26—27 segments (26.8 + 0.5, 5). Without characteristics on underside forewing. Male genitalia (figs. 127, 271, 318, 388). Cap- sule length 266—287 um (2). Tegumen pro- duced into prominent triangular pseuduncus. Gnathos (fig. 318) with central element very short and inconspicuous, with wide truncate tip. Valva (fig. 271) length 206—214 um (2), inner margin approximately straight, but serrate by prominent setal sockets; tip an inconspicuous, pointed, inwards directed process. Aedeagus (fig. 388) 244—266 um (2), without carinae, a simple tube. Female genitalia (figs. 204, 416). T7 with 6—8 short setae in an indistinct row along posterior margin. T8 appearing as a double sclerite: with two lateral patches of scales and 6—7 long setae. Anal papillae with 13—16 setae. Vestibulum smooth, without sclerite or spiculate pouch. Corpus bursae 650—660 um, sparsely covered with small spines or pectinations; signa clearly dissimilar, longest 394—441 um (3), shortest 321—343 um (3), 3.8—3.9 X as long as wide. Ductus spermathecae with 2¥,—3 convolutions. Larva not examined. Biology. Hostplant: Spiraea media Franz Schmidt. Mine (fig. 500). Egg on leaf-underside against midrib, often in axil between midrib and lateral vein. Early mine linear, straight, following a vein, or occasionally leaf margin, filled with brown, dispersed frass; later abruptly widening into wide, irregular blotch, with blackish dis- persed frass. Life history. Probably univoltine. Larvae found in September—October. Adults reared in February—March (probably indoors) and May—June (Szócs, 1968). Distribution (fig. 549). Only known from Slovakia and Matra moun- tains in Hungary. Remarks. This species was discovered by Povolny & Gregor (1952), who described the mine as Stig- mella sp. Later they named it Stigmella spireae Gregor & Povolny, 1955, but still based this name on mines only. This name therefore re- mains a nomen nudum (Code, art. 13a, 16). Later, Gregor & Povolny (1983) redescribed it under the name Ectoedemia spiraeae and desig- nated a neotype. However, since the 1955 name is not available, the last description is to be re- garded as the original species designation and the neotype as holotype. In a collection of Japanese Nepticulidae, at present under study, there is a species reared from Spiraea japonica L. and S. salicifolia L., which is almost unseparable from spiraeae but has a brown hair-pencil instead of a yellowish- white one. Material examined: 5 6,5 2. — Czechoslovakia: 1 3, 1 ©, (paratypes) Slovakia centr. Murän, Huta: Ciganka, 26.1x.1981 on Spiraea media, Gregor & Pov- olny; 1 4, 1 ®, (paratypes), Slovakia or., Slovensky Raj, Cingov, 27.1x.1981 on Spiraea media, Gregor & Povolny (ZMA, EvN). — Hungary: 3 d, 3 2, Matra Hegyseg, Sasto, e.l. 13—19.v.1973, Spiraea media, J. Szöcs (TMAB, ZMA). Mines. — Czechoslovakia: Erzgebirge, Sitno near Banska Stiavnica, Gregor & Povolny (BMNH); Slo- vakia or., Slov. Raj., Cingov, Gregor & Povolny . (ZMA). — Hungary: Matra-Gebirge, Sasto (BMNH). 41. Ectoedemia (Ectoedemia) agrimoniae (Frey, 1858) figs278, 131,152, 2095270) 3195 392746275015 529) Nepticula agrimoniae Frey, 1858: 44, 45. Lectotype d (here designated), Germany: Regensburg, Hof- mann, Frey coll., Genitalia slide 22676 (BMNH) [examined]. Nepticula agrimoniella Herrich-Schaffer, 1860: 60. Syntypes, Germany: Regensburg (Hofmann, An- gerer) (depository unknown) [not examined]. Nepticula agrimoniella; Herrich-Schäffer, [1861]: fig. 169; Heinemann, 1862: 312, 313; Wocke, 1871: 338; 1874: 101; Heinemann & Wocke, 1877: 757, 758; Meyrick, 1895: 722; Sorhagen, 1922: 49, pl. 3 fig. 49; Meyrick, 1928: 859. Nepticula agrimoniae; Ballett Fletcher, 1882: 211; Tutt, 1899: 313—315; Rebel, 1901: 226; Meess, 1910: 479, pl. 91 fig. 68; Petersen, 1930: 68, fig. 92 (4 genitalia). Dechtiria agrimoniae; Beirne, 1945: 205, fig. 61 (6 genitalia). Stigmella agrimoniae; Gerasimov, 1952: 224; Kli- mesch, 1961: 759; Lhomme, 1963: 1192; Borkowski, 1970: 544, figs. 8, 23 (mine, exter- nals). Stigmella (Dechtiria) agrimoniae; Hering, 1957: 41, fig. 19a (mine). Trifurcula (Ectoedemia) 1971: 245. Ectoedemia agrimoniae; Bradley et al., 1972: 2; Borkowski, 1975: 491; Emmet, 1976: 191, pl. 6 He 1, poll, 112 is, 22, Nepticula agrimomella (sic!); Rössler, 1881: 337 [mis- spelling]. agrimoniae; Johansson, Diagnosis: externally similar to species of an- Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 67 gulifasciella complex, but separated by absence of hair-pencil in male, slightly pointed oviposi- tor in female and brown edged scape. Separated from smaller E. hexapetalae, mahalebella and spiraeae by dark collar and edged scape. Both male and female genitalia highly characteristic. Description. Male (fig. 78). Forewing length (1.84) 2.28— 2.96 mm (2.58 + 0.21, 15), wingspan (4) 5.2— | 6.4 mm. Head: frontal tuft yellowish to ferrugi- nous brown, sometimes completely brown; col- lar greyish brown, different from frontal tuft. | Antennae with 35—41 segments (38.3 + 1.7); scape white, but caudal edge with some brown | scales. Thorax and forewings fuscous black with a yellowish silver medial fascia, constricted in middle. Hindwing without hair-pencil or costal bristles. Female. Forewing length 2.0—2.48 mm (2.24 + 0.15, 20), wingspan 4—5.6 mm. Antennae with 31—36 segments (33.2 + 1.3, 15). Thorax and forewings darker than in male, fascia more shining silver. — Male genitalia (figs. 131, 132, 270, 319, 394). | Capsule length 214—240 um (225.7 + 8.4, 6). Tegumen produced into pointed, cuspidate pseuduncus. Gnathos (fig. 319) with triangular, pointed central element, with smooth margins. | Valva (fig. 270) length 163—189 um (174.3 + 10.7, 6), widest at base, distinctly constricted below pointed and inwards curved tip. Aedea- gus (fig. 394) 184—227 um (204.3 + 16.6, 6), dorsal carinae inserted clearly below apex, each divided into 4—5 pointed teeth; ventral projec- tion with some small spines. Female genitalia (figs. 209, 462). T7 with 6—8 | small setae along posterior margin. T8 in form of a narrow curved band, almost split in middle, with a group of scales and 4—7 setae on either side. Anal papillae narrow, with 7—11 setae. Vestibulum without vaginal sclerite, or spiculate pouch. Corpus bursae 440—640 um, complete- ly covered with pectinations; signa similar, cells particularly spiny, length 180—300 um (237.9 + 35.0, 14), 2.3—3.6 X as long as wide. Ductus spermathecae with 3—3Y, convolutions. Larva. Greenish yellow, with conspicuous brown ganglia, head-capsule brown. Without ventral plates. Biology. Hostplants. Agrimonia eupatoria L. and Are- monia agrimonoides (L.) DC. (Greece only). Mine (fig. 501). Egg on leaf-underside. Early mine narrow tortuous gallery, sometimes fol- lowing vein, with broken linear frass, occasion- ally partly contorted; later widening into a wide irregular gallery, or elongate blotch with dis- persed frass. Cocoon made in mine. Life history. Univoltine, larvae from the end of August until October, pupae inside the mine. Adults from May to July. Distribution (fig. 529). Widespread in Central Europe, the Balkans and France, local in South England and south- east Sweden. Not recorded from Denmark, the Netherlands, Belgium, Iberian Peninsula or Italy. Material examined: 49 gd, 67 2, 91 ex.. — Austria: 4 d, 11 ©, Hainburg: Hundsheimer Berg, 200—400 m, el. 15—28.v.1984, Agrimonia eupatoria, J. J. Boomsma & E. J. van Nieukerken (ZMA). — Czechoslovakia: 1 ©, Praha (Prag), Pock. (NMW). — Germany, West: 1 9, Baiern, 1858 (NMW); 1 6,1 9, Frankfurt am Main, coll. Staudinger (MHUB); 1 6, 1 e, Hafen, el. iv.1928, Agrim. eupat., A. Wörz (LNK); 2 d, München, coll. Staudinger; 1 d,2 9, Regensburg, coll. Staudinger (MHUB); 1 d, 1 9, (lecto- and paralectotype of agrimoniae), Regensburg, Hofmann (BMNH); 2 4, 1 ©, Wolfenbuttel, [Heine- mann] (MHUB). — Germany, without further data: 1 ©, Jos. Mann; 1 6, 1 ®, ex coll. v. Heinemann (RMNH); 1 ©, 1869, Lederer (NMW). — Germany, East: 1 6, 1 2, Berlin-Finkenkrug, el. 15— 21.110303 Hlerine; 0 17 eee Berlin Erohnaus sel 10.v.1924, Hering; 91 ex., Berlin, MAJ, Agrimonia, Hering; 5 d, 6 2, Berlin Rudersdf., el. 22.iii— 10.1v.1928, Hering; 7 dg, 7 2, Chorin (Mark), el. 1— 30.1v.1921, Hering (MHUB); 6 d, 6 2, Potsdam, el. 16—22.111.1894, Hinneberg (MHUB, NMW, ZMA); 1 2, [Potsdam] e.l. 16.11.1892, Agrimon. (ZMA). — Great Britain: 3 2, Box Hill, el. 25.vi.1936, 11.v1.1938, 6.v1.1939, S. Jacobs (ZMA); 2 6,1 2, W. of Hadleigh (Essex), South Benfleet, el. 29.vi— 7.v11.1982, G. Bryan & S. B. J. Menken (ZMA); 1 6, 1 ?, no further data, Tyerman, ex coll. BMNH (ZMA). — Greece: 4 4, 2 2, Evvoia: Dhirfis Oros, S. slopes 700—900 m, e.l. 2—18.v.1981, Aremonia agrimo- noides, S. B. J. Menken & E. J. van Nieukerken; 1 4, 3 ©, Frangista (Evritania), valley, 600 m, el. 16.v— 3.vi, 1981, Aremonia and Agrimonia, S. B. J. Menken & E. J. van Nieukerken; 2 d, 6 ©, Katsika (Ioannina) near Limni Ioanninon, 480 m, e.l. 7—15.v.1981, Agrı- monia eupatoria, S. B. J. Menken & E. J. van Nieu- kerken; 2 d, 1 2, Métsovon (loánnina), 950—1000 m, el. 14—22.v.1981, Agrimonia eupatoria, S. B. J. Men- ken & E. J. van Nieukerken (ZMA); 1 d, 2 ©, Vard- housia O., (Fthióus), Dafni, 7 km SE Marmara, 1100 m, e.l. 5—14.v.1981, Agrimonia eupatoria, S. B. J. Menken & E. J. van Nieukerken. — Switzerland: I 6, no further data, 1869 (NMW). — USSR: 3 à, 4 +O 68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Bendery (Tighina), Bessarabia, e.l. 10.1v—20.v.1931, Agrimonia eupatoria, Hering (MHUB). Mines. — On Agrimonia eupatoria. — Austria: Hundsheimer Berg near Hainburg. — Germany: Ber- lin-Frohnau, Hering (BMNH). — Great Britain: Hadleigh; Dorking, Box Hill (Surrey). — Greece: SE Marmara, Vardhoúsia Ori (Fthtiotis). — USSR: Bendery (Tighina), Hering (BMNH). — Yugoslavia: Otoëac. On Aremonia agrimonoides. — Greece: Ev- voia, Dhirfis Oros; SE Marmara, Vardhousia Ori (Fthiötis); Frangísta (Evritania); Fournäs (Evritania). 42. Ectoedemia (Ectoedemia) hexapetalae (Szöcs, 1957) comb. n. (figs. 79, 130, 210, 272, 320, 389, 403, 404, 463, 502, 549) Nepticula utensis Weber var. biol. hexapetalae Szöcs, 1957: 322, 323. Holotype d, Hungary: Budapest, Sashegy, 24.vii.1956 e.l., Szöcs, Genitalia slide 944 Gozmany (TMAB) [examined]. Nepticula hexapetalae; Szöcs, 1965: 79; 1968: 228. Trifurcula hexapetalae; Kasy, 1980: 47. Diagnosis: this species differs externally from most species of the angulifasciella group by its small size, light collar, straight non-metallic fas- cia, and absence of hair-pencil in male. It can possibly be confused with E. mahalebella, in which case the genitalia should be examined. Male genitalia are immediately recognised by the width and the dorsal spinose process of the aedeagus. Female genitalia are easy to separate from mahalebella by shape and position of sig- na. Description. Male. Forewing length 1.96—2.12 mm (2.05 + 0.07, 6), wingspan 4.4—4.7 mm. Head: fron- tal tuft yellowish orange to orange brown; col- lar slightly lighter. Antennae with 30—33 seg- ments (32 + 1.2, 5). Thorax and forewings brownish black with a medial, almost straight fascia, dull white, not shining. Hindwing with- out hair-pencil or costal bristles. Female (fig. 79). Forewing length 1.68—2.04 m (1.89 + 0.13, 8), wingspan 3.7—4.6 mm. Antennae with 24—26 segments (24.9 + 0.8, 8). Male genitalia (figs. 130, 272, 320, 389, 403, 404). Capsule length 197—240 um (3), wider than long. Tegumen distinctly produced into a rounded pseuduncus. Gnathos (fig. 320) divided into short distal element, and basal part with serrate margin. Valva (fig. 272) length 167—184 um (3), relatively broad, inner margin almost straight, but slightly concave below pointed tip. Aedeagus (figs. 389, 403, 404) 261—287 um (3), distinctly longer than capsule, relatively broad; with pair of single or bifid carinae and a single dorsal projection with many spines. Female genitalia (figs. 210, 463). T7 with 4—6 scattered setae along posterior margin. T8 with two lateral patches of scales and 3—5 setae. Anal papillae with 7—18 setae. Vestibulum with incomplete vaginal sclerite with an indistinct ventral projection, without spiculate pouch. Corpus bursae 460—630 um, completely cov- ered by pectinations, especially dense near ves- tibulum; signa similar, with only slight differ- ences in length, 197—326 um (254.7 + 47.5, 7), 2.6—3.1 X as long as wide. Ductus spermathe- cae with 2—3 convolutions. Larva. Pale green, according to Szöcs (1957). Biology. Hostplant. Filipendula vulgaris Moench (= hexapetala Gilibert). Mine (fig. 502). Egg on leaf-underside. Mine © narrow gallery, often following leaf-margin; early mine filled with brown dispersed frass, later black dispersed frass leaving clear margins. Life history. Probably bivoltine. Larvae most abundant in June and July, again in lower num- bers in August and October (Szöcs, 1968). Adults from summer larvae emerged within a month, from autumn larvae in May (only 1 specimen examined). The only specimen taken at light flew in May. Distribution (fig. 549). Still only known from the region near Buda- pest and the Fischawiesen near Gramatneusiedl in the Vienna region. The population of the lat- ter locality appears to be threatened, because these meadows are yearly completely mowed (pers. comm. Kasy), without leaving any old leaves for the autumn generation. Remarks. Originally described as variety of utensis (= angulifasciella) only, but E. hexapetalae appears to be a very distinctive species. Together with terebinthivora these are the only European Ec- toedemia species which are known to be bivol- tıne. Material examined: 6 d,9 2. — Austria: 1 2, Gra- matneusiedl, Fürbachwiesen (= Fischawiesen), e.l. DEAL IB KAS 3 Ls idem, el. 19—20.vu.1979; 1 3, idem, 30.v.1979, at light (NMW). — Hungary: 2 6, 2 2, Budaörs, el. 15.vii.1962, 3.vu.1964, 6— 8.vu.1968, J. Szöcs; 1 6, 1 ©, Budaörs, Törökugrató, el. 29.vi.1968, 21.v.1979, J. Szöcs; 1 d, 1 ©, (Holo- and paratype), Budapest, Sasshegy, el. 24— 26.vu.1956, J. Szöcs (TMAB). Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 69 Mines. — Austria: Gramatneusiedl, Fischawiesen, leg. Kasy. — Hungary: Budapest, Sasshegy, leg. Szöcs (BMNH). The Ectoedemia angulifasciella complex This is a complex of four very similar species, mining on Rosaceae. The status of the four taxa has recently been discussed by Wilkinson et al. (1983), where it was shown that the four taxa form two pairs of sibling species. In that paper the forms schleichiella and staphyleae were not treated, but it is shown here that these are syno- nyms of E. angulifasciella and atricollis respecti- vely. As in the other complexes treated here, the first species (angulifasciella) is described fully, and the other species only as far as they differ from it. 43. Ectoedemia (Ectoedemia) angulifasciella (Stainton, 1849) (es 0332/18 212° 273,321, 399, 464, 503, 532) Nepticula angulifasciella Stainton, 1849: 29. Syntypes, England, Stainton (depository unknown) [not ex- amined]. Nepticula schleichiella Frey, 1870: 286. Lectotype 2 (here designated), Switzerland: Zürich, Frey, Genitalia slide 22567 (BMNH) [examined]. Syn. nov. Nepticula utensis Weber, 1937a: 669, fig. 2. Lectotype 3 (here designated), Switzerland: Zürich, Ute, 28.1x.1935, Sanguis. offic., Z. 2368, Weber, Geni- talia slide ETH 1240 (ETHZ) [examined]. Syn. nov. Nepticula minorella Zimmermann, 1944: 118, 119, figs. 5, 7. Lectotype d (here designated), Austria: Gumpoldskirchen near Wien, e.l. 26.v11.1943, Po- terium min., F. Zimmermann (labelled paratypus). Genitalia slide No. 763/1943 M. Hering (on pin) (MHUB) [examined]. Syn. nov. ? Nepticula brunniella Sauber, 1904, Syntype mines, Germany, West: Hamburg, Sorhagen (depository unknown) [not examined]. Nepticula angulifasciella; Stainton, 1854: 304; Her- rich-Schaffer, 1855: 350; Stainton, 1855: 88—97, pl. 1 fig. 3; Frey, 1857: 417, 418; Stainton, 1859: 435; Heinemann, 1862: 314, 315; Wocke, 1871: 338; 1874: 101; Heinemann & Wocke, 1877: 758, 759; Sorhagen, 1886: 308; Meyrick, 1895: 859; Tutt, 1899: 308—310; Rebel, 1901: 226; Meess, 1910: 479; Meyrick, 1928: 859; Petersen, 1930: 69, fig. 94 (d genitalia); Szócs, 1965: 78. Dechtiria angulifasciella; Beirne, 1945: 205 (partim, not fig. 68); Vari, 1951: 196, 197, figs. 13, 17 (d genitalia, identity). Stigmella angulifasciella; Klimesch, 1951: 62; Gerasi- moy, 1952: 225: Klimesch, 1961: 759; Lhomme, 1963: 1193; Borkowski, 1969: 112. Stigmella (Dechtiria) angulifasciella; Hering, 1957: 902 (mine). Trifurcula (Ectoedemia) angulifasciella; Johansson, 1971: 245. Ectoedemia angulifasciella; Bradley et al., 1972: 2; Emmet, 1973: 178—180 (differences with atricol- lis); 1976: 192, pl. 6 fig. 2, pl. 12 fig. 24; Wilkinson et al., 1983: 211—224, figs. 1, 2, 9 (specific status). Ectoedemia angulifasciella (partim); Borkowski, 1975: 492. -Trifurcula angulifasciella (partim); Karsholt & Niel- sen, 1976: 18. Nepticula schleichiella; Wocke, 1871: 338; Heine- mann & Wocke, 1877: 759, 760; Rebel, 1901: 226; Meess, 1910: 479. Stigmella schleichiella; Gerasimov, 1952: 259; Hering, 1957: 937 (mine). Stigmella utensis; Klimesch, 1948: 72, 73, figs. 50, 51 (4 genitalia). Stigmella minorella; Klimesch, 1961: 739. ? Nepticula brunniella; Sorhagen, 1922: 59, fig. 70. Diagnosis: male characterised by the combi- nation of a yellowish-orange collar, an oblique metallic fascia and a white hair-pencil. E. spi- raeae is very similar, but has almost no metallic fascia and is usually smaller. Male genitalia characterised by the shape of the valva, with si- nuous inner margin. Female separated from ag- rimoniae, atricollis and arcuatella by light col- lar, E. mahalabella is very similar, but usually smaller and with very different signa. Description. Male (fig. 80). Forewing length (excluding specimens reared from Filipendula) 2.2—2.8 mm (2.56 + 0,18, 21), wingspan 5.2—6.6 mm. Including Fihpendula specimens: forewing length 1.92—2.8 (2.47 + 0.25, 25), wingspan 4.46.6 mm. Head: frontal tuft and collar pale ochreous to ferruginous, usually lighter than in atricollis; collar often slightly lighter. Antennae with 29—35 segments (32.2 + 1.6, 18). Thorax and forewings fuscous black, with a medial, oblique, shining metallic silver fascia, rarely in- terrupted in middle. Underside of forewing with small scaleless area. Hindwing with white hair-pencil and a few dark scales along costa. Female. Forewing length (excluding speci- mens reared from Filipendula) 2.04—2.68 mm (2.52 + 0,18, 12), wingspan 4.7—6.1 mm. In- cluding Filipendula specimens: forewing length 1.92—2.68 (2.40 + 0.26, 16), wingspan 4.5—6.1 mm. Antennae with 25—29 segments (27.1 + 1.0, 13). Male genitalia (figs. 133, 273, 321, 390). Cap- sule length 210—257 um (241.0 + 16.6, 9). Te- 70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 gumen distinctly produced into truncate pseud- uncus. Gnathos (fig. 321) with central element divided, distal part spatulate, basal part with serrate margin. Valva (fig. 273) length 159—193 um (175.3 + 11.6, 11), inner margin sinuous, forming a slight, but distinct rounded bulge in distal half, so that inner margin forms a right angle with pointed tip. Aedeagus (fig. 390) 214—274 um (250.1 + 18.3, 11), slightly con- stricted beyond middle, carinae single or bifid, not sharply pointed, with many small spines at base. Female genitalia (figs. 211, 212, 464). T7 without setae in a row. T8 with two lateral patches with many scales and about 4—8 setae; anal papillae with 5—9 setae. Vestibulum with a vaginal sclerite and a “spiculate” pouch without spines. Corpus bursae 400—570 um, almost completely covered with pectinations; signa dis- similar, longest 249—381 um (326.1 + 38.0, 10), shortest 227—356 um (289.5 + 37.1, 9), 3.3— 4.6 X as long as wide. Ductus spermathecae with 2/—3 convolutions. Larva. Greenish white, with distinct ganglia. Head-capsule and prothoracic plate dark brown. In 2nd and 3rd instar with chain of dark brown ventral plates. Biology. Hostplants: Rosa spp., including evergreen Rosa sempervirens L., occasionally on Sangui- sorba minor Scop., S. officinalis L. and Filipen- dula vulgaris Moench (in Hungary only). Mine (fig. 503). Egg on leaf-underside. Early mine highly contorted gallery filled with brown, contorted frass; later widening into large irregu- lar blotch or wide gallery with irregular dispers- ed black frass. Life history. Univoltine. Larvae from end of August to early November. S. E. Whitebread (in litt.) found some larvae in July in Switzer- land. Adults flying from the middle of June to the end of July. May records probably refer all to indoor rearing. Distribution (fig. 532). Widespread in Europe, from southern Scan- dinavia to Greece. Not yet recorded from Ire- land, Iberian Peninsula and central Balkan. Remarks. In the Stainton collection there are only an- gulifasciella specimens collected after 1849, thus without syntype status. The type specimens were not reared, but were later recognised by Stainton as being the same species as the rose miner. The identity of this species has been dis- cussed by Wilkinson et al. (1983), with excep- tion of the Sanguisorba form. Three authors de- scribed the Sanguisorba form: Frey as schlei- chiella, Weber as utensis (from the same locality as Frey!) and Zimmermann as minorella. The lectotypes of these taxa are morphologically identical with angulifasciella, and also the biolo- gy, except the foodplant, is similar. Electropho- resis of one larva collected in the Pyrenees on Sanguisorba minor showed that this form is also genetically identical with angulifasciella (Men- ken, in preparation). The conclusion is that an- gulifasciella ıs-an oligophagous species, which most commonly feeds on Rosa. Sz6cs also col- lected the species in numbers on Filipendula vulgaris in Hungary. These specimens are much smaller than normal angulifasciella probably . due to the size of the leaves. The measurements of the adults have thus been given both exclud- ing and including these specimens. N. brunniella Sauber has been described on the basis of some mines collected by Sorhagen in Hamburg. Judging from Sorhagen’s (1922) description and figure they could also belong to angulıfasciella. Material examined: 38 d, 28 ©, 42 ex. — Austria: 3 3, (lecto- and paralectotypes of minorella), Gumpoldskirchen near Wien, el. 14.vi—26.v11.1943, Poterium min., Zimmermann (MHUB); 1 4, Gumpoldskirchen, Glaslauterriegel, 28.v11.1972, F. Kasy; 2 6, Hundsheimer Berg (near Hainburg), (Evil 77 5151980 NE Ma sy ENE ZVL Knitsche (NMW). — France: 1 6, 1 2, Chaville, e.l. 31.v., Joannis (MNHN). — Germany, West: 1 d, Bayers, 1858 (NMW); 2 6, 1.5 km NW Birresborn (Rhl.-Pf.), Vulkanberg, 460 m, e.l. 24—27.v1.1983, Rosa, Alders & Van Nieukerken (ZMA); 2 ®, Braunschweig, Heinemann, coll. Staudinger (MHUB); 2 d, 1 9, Stuttgart, 30.v1.1883; 19.vu.1886 (MHUB, NMW); 1 &, no data, ex coll. Heinemann, coll. Snellen (RMNH). — Germany, East: 1 d, 1 9, Friedland, 1, 9.v.1885, Stange; 5 d, 3 ©, Rachlau, el. 1897, Rosa canina, Schütze; 2 ©, Sachsenberg, Nord- hausen, e.l. 29.vi.1899, Rosa, Petry (MHUB). — Hungary: 5 d, 4 ©, Szär, el. 17—24.v1.1968, Filipen- dula vulgaris, J. Szöcs (TMAB). — Netherlands: 3 6, Nunspeet, el. 14—22.v11.1946, Rosa, L. Vari; 8 d, 5 2, Ootmarsum, Achter de Voort, e.l. 10—14.v11.1981, Rosa, 15.x.1980, Andeweg & Van Nieukerken; 3 ®, Winterswijk, quarry, e.l. 26.v11.1979, 12—13.v11.1982, Van Nieukerken (ZMA). — Poland: 42 ex., Krosno Odr. (Crossen a. Oder), e.l. 10—26.v1.1930, Rosa canina, Hering; 3 d, 3 ©, idem, el. 15—23.v.1932; 1 3, Silesia, Wocke (MHUB); 1 à, Silesia, 1872, Stau- dinger (NMW); 2 6, 4 ©, Wroclaw (Breslau), el. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 71 iv.1872, Rosa (Wocke) (MHUB). — Switzerland: 2 ? (lecto- and paralectotype of schleichiella), Zürich, Frey (BMNH); 1 d (lectotype of utensis, see above); 2 4,1 2, Zürich, Uto, mine 29.ix.1936, Sanguisorba officinalis, Weber (ETHZ). — Yugoslavia: 3 d,2 9, Selce, 4 km SE Crikvenica (Hrvatska), a.s.l., e.l. 24.v—18.vi.1984, J. J. Boomsma & E. J. van Nieuker- ken (ZMA). — No data: 1 6, Rosa (ZMA); 2 9, el. 6.vi.1884, v. 1903, Rosa (NMW). Mines. — On Rosa spp. — Austria: Gumpoldskir- chen. — France: Andlau; Arvieu; Barr; Corse, Portic- cio; Modane. — Germany, West: Alendorf; Birres- born; Hillesheim; Klotten. — Great Britain: Saffron Walden; Takeley; Tintern. — Greece: Fournas, Evri- tania; Oiti Oros, Fókis + Fthiótis; Olympia; Parnos Oros, Attika. — Italy: Picinisco; Trento. — Nether- lands: Cadier en Keer; Epen; Kunrade; De Lutte; Ootmarsum; Winterswijk; Wijlre. On Sanguisorba minor. — France: Porté-Puymorens. — Germany, West: Alendorf. 44. Ectoedemia (Ectoedemia) atricollis (Stainton, 1857) (figs. 15—17, 28, 81, 134, 213, 214, 274, 323, 391, 465, 504, 505, 533) Nepticula atricollis Stainton, 1857: 112. Lectotype © (here designated), England, ex Boyd Coll. B.M., 1813—391, 2 5788, Nepticula atricollis Stn. Type, Genitalia slide 22617 (BMNH) [examined]. Nepticula atricolella Doubleday, 1859: 36 (unjustified emendation). Nepticula aterrima Wocke, 1865: 270. Lectotype d (here designated), Poland: Freiburg, Silesia, e.l. iv.1862, Crataegus, Wocke, Genitalia slide VU 2325 (ZIAS) [examined]. Nepticula malivora Toll, 1934b: 70, 83, pl. 2 fig. 1. Nomen nudum (no description or diagnosis, mine only). Nepticula atricollis var. aterrimoides Skala, 1940: 143. Nomen nudum (no description or diagnosis). Nepticula staphyleae Zimmermann, 1944: 117, 118, figs. 4, 6. Lectotype d (here designated), Austria: Gumpoldskirchen near Wien, e.l. 12.v1.1943, Sta- phylea pinnata, F. Zimmermann, Genitalia slide VU 1488 (MHUB) [examined]. Syn. nov. Nepticula atricollis var. prunivora Skala 1941: 1977. Nomen nudum (no description or diagnosis, mine only). Nepticula atricollis; Stainton, 1859: 435; 1862: 228— 25 D AE ls K-leinemann, 1862: 313, 314; Nolcken, 1871: 782; Wocke, 1871: 338; 1874: 101; Heinemann & Wocke, 1877: 758; Meyrick, 1895: 722; Tutt, 1899: 304—306; Rebel, 1901: 226; Meess, 1910: 479; Meyrick, 1928: 859; Peter- sen, 1930: 69, fig. 93 (3 genitalia); Klimesch, 1936: 208; Zimmerman, 1944: fig. 6 a—c (d geni- talia); Szöcs, 1965: 79. Dechtiria atricollis; Vari, 1951: 197 (comparison with angulifasciella); Emmet, 1971: 171, 240, 241. Stigmella atricollis; Gerasimov, 1952: 228, Klimesch, 1961: 759; Lhomme, 1969: 104. Stigmella (Dechtiria) atricollis; Hering, 1957: 349, 664, 690, 835, 854, 1010; figs. 229b, 408a. Trifurcula (Ectoedemia) atricollis; Johansson, 1971: 245. Ectoedemia atricollis; Bradley et al., 1972: 2; Emmet, 1973: 178—180 (differences with angulifasciella); Emmet, 1976: 193, pl. 6 figs. 4, 5, pl. 12 fig. 25; Wilkinson et al., 1983: 211—224, figs. 3, 4, 10 (specific status). Dechtiria angulifasciella (partim); Beirne, 1945: 205, fig. 68 (4 genitalia). Ectoedemia angulıfasciella 1975: 492. Trifurcula angulifasciella (partim); Karsholt & Niel- sen, 1976: 18. Nepticula aterrima; Wocke, 1871: 338, 1874: 102; Heinemann & Wocke, 1877: 763; Rebel, 1901: 227; Meess, 1910: 480. Stigmella aterrima; Gerasimov, 1952: 228; Lhomme, 1963: 1198. Nepticula malivora; Toll, 1936: 411. Nepticula staphyleae; Szöcs, 1965: 79. Stigmella staphyleae; Hering, 1957: 1027 (mine); Kli- mesch, 1961: 759. Ectoedemia staphyleae; Borkowski, 1975: 493. 1963: 1193; Borkowski, (partim); Borkowski, Diagnosis: separated from angulıfascıella, mahalebella and spiraeae by dark collar, from agrimoniae by hair-pencil in male and blunt ovipositor in female (pointed in agrimoniae), from rubivora by head colour and from spino- sella by size, and dark coloured hair-pencil in male spinosella. E. arcuatella can hardly be dis- tinguished from atricollis, except by smaller size, much shorter signa and shorter aedeagus of arcuatella. Description. Male. Forewing length 2.16—2.56 mm (2.39 + 0.12, 20), wingspan 4.8—6.0 mm. Head: frontal tuft orange to ferruginous (rarely black); collar dark fuscous to black. Antennae with 29—39 segments (33.3 + 2.4, 16). Hindwing with white hair-pencil, surrounded by some dark brown scales, especially along costa. Fur- ther as angulifasciella. Female (fig. 81). Forewing length 2.28—2.80 mm (2.56 + 0.14, 14), wingspan 5.2—6.2 mm. Antennae with 26—30 segments (27.7 + 1.2, 12). Male genitalia (figs. 134, 274, 323, 391). Cap- sule length 270—287 um (278.6 + 6.9, 11). Gnathos fig. 322. Valva (fig. 274) length 176— 206 um (189.4 + 8.5, 11), inner margin almost straight, forming an obtuse angle with pointed 72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 tip. Aedeagus (fig. 391) 261—287 um (273.1 + 8.8, 11), hardly constricted. Further as angulı- fasciella. Female genitalia (figs. 213, 214, 465). T8 with 3—4 setae at both sides. Anal papillae with 6—9 setae. Spiculate pouch with few almost invisible spines. Corpus bursae 495—660 um; longest signum (360) 411—489 um (435.5 + 32.3, 12), shortest (356) 377—446 um (405 + 24.4, 12), 3.8—5.0 X as long as wide. Ductus spermathe- cae with 3—3'/, convolutions. Larva. Greenish white, with distinct ganglia. Head-capsule and prothoracic plate black. In 2nd and 3rd instar with chain of black ventral plates. Biology. Hostplants. Oligophagous on Rosaceous trees: most abundant on Crataegus spp., com- mon on Malus sylvestris Miller, Pyrus communis L. and Prunus avium L., occasionally on Prunus mahaleb L. and P. cerasifera Ehrh. Records on Prunus spinosa L. probably all refer to E. spino- sella. In east Central Europe also common on Staphylea pinnata L. (Staphyleaceae). Mine (figs. 504, 505). Egg on leaf-underside. Early mine linear, following leaf-margin, or slightly contorted, filled with brown frass; later widening into large blotch with scattered black frass. Life history. Univoltine. Larvae from late August until late October, commonest in Sep- tember. Adults slightly earlier than angulifas- ciella, from early June until late July. May re- cords probably refer to indoor rearing. Distribution (fig. 533). Widespread in Europe, from Central Sweden to Central Italy. Not yet found in a large part of the mediterranean region and Ireland. Remarks. Beirne (1945) and Borkowski (1975) synony- mised this species with angulifasciella, but Wil- kinson et al. (1983) showed that both species are separate, genetically isolated entities. N. aterrıma Wocke is just a dark aberration of atricollis. The nomina nuda malivora Toll and aterrimoides Skala are based on mines of atricollis. N. staphyleae Zimmermann is morphologi- cally identical to atricollis, the adult, larva, and mine being completely similar. The hostplant of staphyleae is however unrelated to the Rosa- ceae. By analysis of allozymes (Menken, in preparation) the larvae collected from Staphylea in the autumn of 1983 are shown to be geneti- cally identical to those of sympatric atricollis from Crataegus. Therefore staphyleae is here synonymised with atricollis. Material examined: 64 6, 59 2, 1 ex. — Austria: 3 3, 1 2 (lecto- and paralectotypes of staphyleae), Gumpoldskirchen near Wien, el. 12.vi—21.vii.1943, Staphylea pinata, Zimmermann (MHUB); 1 4,1 2, 1 km N. Gumpoldskirchen, Richardshof, e.l. 5.vi.1984, Staphylea pinnata, J. J. Boomsma & E. J. van Nieu- kerken (ZMA); 1 9, Klosterneuburg, Buchberg, e.l. 24.v.1937, Preissecker; 1 2, Linz, 9.11.1911, Knitsche; 1 3, Wien, Haschbg., el. 22.v.1937, Preissecker (NMW). — Czechoslovakia: 1 6, 1 ©, Decin (Tets- chen, Elbe), el. 11, 23.v1.1943, Crataegus, Hering (MHUB). — France, 2 6, 3 2, Clamart (Hauts de Seine), e.l. 4.vi, Aubépine (Crataegus), De Joannis (MNHN). —- Germany, West: 3 d, 2 9, Freiburg, el. 1v.1965, Pyr. mal., 1 ©, Hannover, Glitz; 2 ©, Wolf- . enbuttel, [Heinemann] (MHUB). — Germany, East: 10 3, 9 ©, Berlin-Finkenkrug, el. 31.v—10.vi.1930, Pyrus malus, Hering; 2 4, 4 2, Bredow b. Nauen, e.l. 25.1—2.1v.1925, Malus silvestris, Hering; 2 6, 2 9, Rüdingsdorf, Nordhausen, el. 21.v—7.vi.1921, 1.v1.1925, Crataegus, Petry; 2 d, 4 2, Rachlau, el. 1.1888, 1897, Pyrus malus, Schutze (MHUB). — Great Britain: 1 2 (lectotype, see above). — Hungary: 2 6, Budapest, e.l. 16.vi.1953, Staphylea, J. Szöcs; 1 9, Budapest, Csittepéta, e.l. 30.v.1978, Staphylea pinna- ta, J. Szöcs; 1 2, Normafa, e.l. 14.v.1978, Staphylea pinnata, J. Szöcs; 1 9, Budapest, Zugliget, el. 23.vi.1957, Staphylea pinnata, J. Szöcs (TMAB). — Italy: 1 2, Formello (Roma), Valle delle mad. d. Sor- bo, e.l. 5.vi.1984, Crataegus monogyna, S. B. J. Men- ken (ZMA). — Netherlands: 32 d, 20 2, from fol- lowing localities; Ankeveense Plassen, Castricum, Loosdrecht, Nederhorst den Berg, St. Pietersberg, Weesp, Winterswijk and own breeding, reared from Crataegus, Malus or Pyrus (RMNH, ZMA). — Po- land: 1 d (lectotype of aterrima, see above); 2 4,19, Silesia, Wocke, Staudinger (MHUB, ZMA). — Switzerland: 1 d,1 2, 1869, 1870 (NMW). Mines. On Crataegus. — Austria: Hundsheimer Berg near Hainburg; Orth am Donau. — Germany, West: Bad Honnef; Birresborn; Gerolstein. — Great Britain: Chepstow; Churchill; New Forest; Takeley. — Netherlands: many localities. — Italy: Formello; Opi. On Malus. — Austria: Orth am Donau. — Great Britain: Stapleford Abbots. — Italy: Picinisco. — Netherlands: Denekamp; Hilversum; Leiden; Neder- horst ten Berg; Rockanje; Wassenaar; Winterswijk. On Mespilus germanica. — Netherlands: Winters- wijk. On Prunus avium. — Austria: Hof am Leitha- gebirge. — Germany, West: Bad Honnef. — Nether- lands: Oud Valkenburg; Rijckholt; Sibbe; St. Geer- truid; Winterswijk. On Prunus cerasifera. — Rumania: Cocos, Niculitel, Tulcea, 1.1x.1973, leg. Draghia. On Prunus mahaleb. — Germany, West: Klotten. Pyrus. — Italy: Opi. — Netherlands: Hilver- | | Nepticula arcuatella Herrich-Schäffer, 1855: | Stigmella (Dechtiria) arcuatella; VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 73 sum; Leiden; Wassenaar; Winterswijk. — Yugoslavia: Slavonska Pozega. On Staphylea pinnata. — Austria: Gumpoldskirchen; Hundsheimer Berg near Hain- burg. — Yugoslavia: N. Bihac. 45. Ectoedemia (Ectoedemia) arcuatella (Herrich-Schäffer, 1855) (figs. 82, 136, 215, 275, 323, 392, 466, 506, 534) 354. Lectotype d (here designated) identical with lec- totype of N. arcuata Frey, see below. Nepticula arcuata Frey, 1856: 384, 385. Lectotype d (here designated), Switzerland: Zurich, Frey, Genitalia slide 22678 (BMNH) [examined]. Nepticula arcuosella Doubleday, 1859: 36 (unjustified emendation). Nepticula arcuata; Frey, 1857: 415—417; Stainton, 1858: 97; 1859: 434, 435; 1862: 196—203, pl. 9 fig. 3 (biology); Nolcken, 1871: 784—786. | Nepticula arcuatella; Heinemann, 1862: 315, 316; Wocke, 1871: 338; 1874: 101; Heinemann & Wocke, 1877: 759; Meyrick, 1895: 723; Tutt, 1899: 306—308; Rebel, 1901: 226; Meess, 1910: 479; Meyrick, 1928: 860; Petersen, 1930: 70, fig. 96 (3 genitalia); Klimesch, 1936: 208; Szócs, 1965: 78. Dechtiria arcuatella; Beirne, 1945: 206, fig. 70 (6 genitalia). Stigmella arcuatella; Klimesch, 1951: Gerasimov, 1952: 226; Klimesch, 1961: 759; Lhomme, 1963: 1194; Borkowski, 1969: 105, figs. 13, 14. Hering, 1957: 42, 454, 821, fig. 503c (mine). Trifurcula (Ectoedemia) arcuatella; Johansson, 1971: 245. Ectoedemia arcuatella; Bradley et al., 1972: 2; Em- met, 1973: 180, 278 (differences with rubivora); Borkowski, 1975: 492; Emmet, 1976: 194, pl. 6 fig. 3, pl. 12 fig. 26; van Nieukerken, 1982: 108; Wilkinson et al., 1983: 211—224, figs. 5, 6, 11 (specific status). Trifurcula arcuatella (partim); Karsholt & Nielsen, 1976: 18. Diagnosis: when not reared almost insepara- ble from E. atricollis, see diagnosis for that spe- cies. Females difficult to separate from spinosel- la. Description. Male (fig. 82). Forewing length 1.80—2.24 m (2.11 + 0.17, 8), wingspan 4.0—4.9 mm. | Head: frontal tuft yellow to ferruginous, mixed with fuscous scales, getting darker towards col- lar; collar fuscous to black. Antennae with 28— | 32 segments (29.9 + 1.5, 8). Hindwing with a white hair-pencil. Further as angulifasciella. Female. Forewing length 1.64—2.32 mm (2.05 + 0.26, 10), wingspan 3.6—5.2 mm. An- tennae with 24—28 segments (26.2 + 1.4, 9). Male genitalia (figs. 136, 275, 323, 392). Cap- sule length 249—253 um (3). Gnathos fig. 323. Valva (fig. 275) length 180—189 um (4), inner margin almost straight, forming an obtuse angle with pointed tip. Aedeagus (fig. 392) 231—244 um (4), hardly constricted. Female genitalia (figs. 215, 466). T8 with about 5 setae at each side. Anal papillae with 5—9 setae. Spiculate pouch with very few min- ute spines. Corpus bursae 420—500 um; longest signum 227—313 um (4), shortest 206—283 um (4), 3.1—4.1 X as long as wide. Ductus sperma- thecae with 2!/, convolutions. Larva. Pale yellow, ganglia not very distinct. Head-capsule and prothoracic plate light brown. Penultimate instars with chain of brown ventral plates, which are shed in final instar. Biology. Hostplants. Fragaria vesca L., F. moschata Duchesne, Potentilla erecta (L.) Räuschel, P. sterilis (L.) Garcke. Mine (fig. 506). Egg on leaf-underside. Early mine highly contorted gallery with brown, coiled frass; later widening into large irregular blotch with scattered brown frass. Life history. Univoltine. Larvae from late August to middle of October. Adults emerge from end of May to July. Distribution (fig. 534). Widespread in Europe, but scarcer than the other three species of the complex. Only one re- cord each from the Netherlands and France. Not yet recorded from Norway, Iberian Penin- sula, Belgium or Ireland. Remarks. Frey discovered this species, named it arcuata and described it in 1856. However, Herrich- Schaffer, who renamed it arcuatella and attrib- uted the species to Frey, described it one year ahead, and therefore is attributed with the authorship. Since Herrich-Schaffer clearly re- fers to the Frey material, it can be regarded as type material for both arcuatella and arcuata. The synonymy of rubivora with this species, as suggested by Borkowski (1975) has been refut- ed by Wilkinson et al. (1983). Material examined: 29 d, 27 2. — Austria: 8 d, 6 2,5 km. W. Völkermarkt: Pörtschach (Kärnten), 500 m, e.l. 19.v—5.v1.1984, Fragaria vesca, J. J. Boomsma & E. J. van Nieukerken (ZMA); 1 6, 12, Wien, 74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Haschbg., e.l. 13, 20.v.1937, Preissecker (NMW). — Denmark: 2 ©, Bornholm, Gudhjem, el. 31.v— Av Eragon (EIS Larsen (ZMO)Z Ger Heinemann many, West: 1 ©, Braunschweig, (RMNH); 2 dg, Freiburg, 11.1882, Fragaria; 1 6, Wolfenbuttel, [Heinemann] (MHUB); 2 ®, Pfalz, Eppelsheim (MHUB, NMW); 1 2, no data, 1870, Heinemann (MHUB); 1 4, locality illegible, Frag. vesc., Heinemann (RMNH); 1 2, no data, 1878, Stau- dinger (NMW). — Germany, East: 1 d, Friedland, 11.iv.1889, Stange (NMW); 1 ©, Kyffhausen, 12.vi.1912 (NMW); 2 ©, Rachlau, Schütze (MHUB). — Greece: 1 d, Frangista (Evritania), valley, 600 m, st. 29, e.l. 13—15.vi.1981, Fragaria vesca, Menken & Van Nieukerken; 1 6, 1 2, 3 km SE Neráidha (Evri- tania), 1200 m, st. 37, e.l. 3—4.vi.1981, Fragaria ves- ca, Menken & Van Nieukerken (ZMA). — Nether- lands: 1 6, 2 2, Woods W. of Wijlre, el. 3— 9.vii.1982, E. J. van Nieukerken (ZMA). — Poland: 1 2, Wroclaw (Breslau), el. iv.1864, Fragarıa (MHUB). — Switzerland: 1 4, 1 © (lecto- and para- lectotype), Zurich, Frey (BMNH). — USSR: 4 6, 2 2, Bendery (Tighina), Bessarabia, e.l. 16—18.v.1931, Fragaria vesca, Hering (MHUB). — Yugoslavia: 1 ?, 2 km W. Otocac (Hrvatska), 450—500 m, e.l. 26— 28.v.1984, Fragaria vesca, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. — On Fragaria vesca. — Austria: Hof am Leithagebirge; Volkermarkt. — Great Britain: Chur- chill; Grays; Saffron Walden; Tintern. — Greece: Fournäs, Evritania; Frangista, Evritania; Neraidha, Evritania. — Italy: Tolmezzo. — Netherlands: Wijlre. — Yugoslavia: Han Knezica, N. of Prijedor; Otocac; Mt. Slavnik, S. of Herpelje-Kozina. On Po- tentilla erecta. — Italy: Tramonti di Sopra. 46. Ectoedemia (Ectoedemia) rubivora (Wocke, 1860) (figs. 83, 135, 216, 276, 324, 393, 413, 467, 507, 535) Nepticula rubivora Wocke, 1860, 132. Syntypes, Po- land: Wroclaw (Breslau), e.l. ıv.18.. (ante 1860), Wocke (depository unknown) [not examined]. Nepticula rubivora; Heinemann, 1862: 315; Nolcken, 1871: 783; Wocke, 1871: 338; 1874: 101; Heine- mann & Wocke, 1877: 783; Meyrick, 1895: 722, 723; Tutt, 1899: 310—313; Rebel, 1901: 226; Meess, 1910: 479; Sorhagen, 1922: 49, 50, pl. 3 fig. 52; Meyrick, 1928: 860; Petersen, 1930: 69, fig. 95 (4 genitalia); Klimesch, 1936: 208; Szöcs, 1965: 76. Dechtiria rubivora; Beirne, 1945: 205, fig. 69 (G geni- talia). Stigmella rubivora; Klimesch, 1951: 62; Gerasimov, 1952: 257; Klimesch, 1961; 759; Lhomme, 1963: 1194; Borkowski, 1969: 112. Stigmella (Dechtiria) rubivora; Hering, 1957: 908, fig. 579a. Trifurcula (Ectoedemia) rubivora; Johansson, 1971: 245. Ectoedemia rubivora; Bradley et al., 1972: 2; Emmet, 1973: 180, 278 (differences with arcuatella); 1976: 195, pl. 6 fig. 7, pl. 12 fig. 27; Wilkinson et al., 1983: 211—224, figs. 7, 8, 12 (specific status). Ectoedemia arcuatella rubivora; Borkowski, 1975: 492. Trifurcula arcuatella (partim); Karsholt & Nielsen, 1976: 18. Diagnosis: separated from the other Rosaceae mining Ectoedemia species by the black head in both sexes. In genitalia almost inseparable from arcuatella, although signa seem to have fewer cells. Description. Male (fig. 83). Forewing length 2.0—2.56 mm (2.28 + 0.12, 25), wingspan 4.6—5.7 mm. Head: frontal tuft and collar black, sometimes with some fuscous scales. Antennae with 30— | 37 segments (33.2 + 1.7, 19). Hindwing with white hair-pencil. Further as angulifasciella. Female. Forewing length 2.08—2.69 (2.43 + 0.18, 28), wingspan 4.6—6.0 mm. Antennae with 25—31 segments (27.9 + 1.6, 24). Male genitalia (figs. 135, 276, 324, 393, 413). Capsule length 257—283 um (269.1 + 9.8, 5). Gnathos fig. 324. Valva (fig. 276) length 176— 206 um (196.3 + 11.9, 5), inner margin almost straight, forming an obtuse angle with pointed tip. Aedeagus (fig. 393) 236—266 um (248.6 + 12.5, 5), hardly constricted. Further as anguli- fascıella. Female genitalia (figs. 216, 467). T8 with few setae at both sides. Anal papillae with 4—6 se- tae. Spiculate pouch with some almost invisible spines. Corpus bursae 410—460 um; longest signum 227—274 um (245.5 + 17.1, 7), shortest 201—257 um (226.5 + 22.6, 7), 2.9—3.7 X as long as wide. Ductus spermathecae with 2Y, convolutions. Larva. Pale yellow, or yellowish white with green tinge, ganglia conspicuous. Head-capsule and prothoracic plate brown. Penultimate instars with chain of dark brown ventral plates and smaller, similar dorsal plates, which are shed in final instar. Biology. Hostplants. Rubus fruticosus L. (sensu lato), R. caesius L., R. saxatilis L., R. chamaemorus L. and R. arcticus L. (Kyrki & Tabell, 1984). Not found on R. idaeus L. Mine (fig. 507). Egg on leaf-underside. Early mine highly contorted gallery filled with brown VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 75 frass; later widening into large irregular blotch with scattered black frass. Often staining sur- rounding tissue purple. Life history. Univoltine. Larvae from late August until late October. Adults fly in June and July. Distribution (fig. 535). Widespread in Europe, from Lapland south- wards to Central Italy. In the mediterranean re- gion usually in river valleys and mountains on- ly. Remarks. According to R. Puplesis (in litt.) no type material of this species is present in Wocke’s collection in Leningrad, but from Wocke’s very clear description and from subsequent Wocke material there can be no doubt about the identi- ty of his species. Wilkinson et al. (1983) dis- cussed the separate identity of rubivora and ar- cuatella. Material examined: 62 6, 74 2. — Austria: 1 6, Linz, Au, 23.v.1923, Knitsche (NMW). — Denmark: 2 ©, Faaborg (Fynen), Alliskus, el. 7—15.vi.1926, Rubus; 1 3, 4 2, Faaborg (Fynen), Sändarsjöen, e.l. 6—10.v1.1920, 17.v1.1922, 12—15.vi.1926, Rubus (ZMC). — Germany, West: 1 d, 2 9, Braunschweig, Heinemann (MHUB); 1 6, Hannover, Lederer (NMW); 1 &, Wolfenbuttel, [Heinemann] (MHUB). — Germany, East: 1 d, 5 ©, Berlin-Finkenkrug, e.l. 6—12.vi.1930, Rubus caesius, Hering; 2 6, 2 9, Friedland, el. 4—10.1v.1888, Rubus caesius, Stange (MHUB); 2 d, idem, iv.1900 (NMW). — Great Bri- tain: 1 d, 1 ©, Saffron Walden (Essex), 3 km NE, el. 24.vi-4.v11.1980, Bryan, Emmet & Van Nieukerken (ZMA). — Italy: 2 d, 1 ©, 4 km WSW Tolmezzo (Udine), Villa Verzegnis, 550 m, el. 16—18.vi.1984, J. J. Boomsma & E. J. van Nieukerken (ZMA). — Netherlands: 43 d, 41 ® from following localities: Blaricum, Gronsveld, Hulshorst, Kortenhoef, Lunte- ren, Nunspeet, Simpelveld, Winterswyk (RMNH, ZMA). — Poland: 1 g, Silesia, Staudinger (RMNH); 1 2, Wroclaw (Breslau), e.l. iv.1869, Rubus caesius (RMNH); 1 4, 5 2, Wroclaw (Breslau), v.1862, [Wocke], Rubus caesius (MHUB). — Switzerland: 2 2, Glarus, el. 20.v., 7.v1.1875, Rubus petraeus (= saxatilis) (MHUB); 2 6, Zurich, coll. Lederer (MHUB); 1 ©, no data, 1868 (NMW). — USSR: 1 d, Estonia, Nomme, Moor, Rub. cham., Petersen (MHUB). — Yugoslavia: 2 2, Mt. Slavnik, 5 km S. Herpelje-Kozina (Slovenia), 800 m, el. 26.v— 7.v1.1984, J. J. Boomsma & E. J. van Nieukerken; 2 2, Sovinjak, 9 km NE Motovun (Hrvatska), Mirna valley, el. 8—15.vi.1984, J. J. Boomsma & E. J. van Nieukerken (ZMA). — No Data: 1 8, el. vi, Rubus caes. (RMNH). Mines. — Austria: Wien, Lobau. — Belgium: Zold- er. — Germany, West: Gerolstein; Oberstadtfeld. — Great Britain: Cheddar Gorge; Grays; Hadleigh; Sat- fron Walden. — Italy: Tramonti di Sopra; Trento; Tolmezzo. — Netherlands: many localities. — Yugoslavia: NE Bihac; S. of Novska; Mt. Slavnik, S. of Herpelje-Kozina; Sovinjak, NE Motovun. 47. Ectoedemia (Ectoedemia) spinosella (de Joannis, 1908) (figs. 18—20, 84, 137, 138, 217, 218, 277, 325, 395, 468, 508, 509, 536) Nepticula spinosella J. de Joannis, 1908a: 328. Lecto- type ® (here designated), France: Vannes, 18.vi., prunetier, L. de Joannis, Genitalia slide VU 947 (MNHN) [examined]. Nepticula spinosella; J. de Joannis, 1908b: 825, 826, fig. 3, pl. 15 fig. 13 (larva, mine, adult); Klimesch, 1936: 206; 1941: 163, 164, pl. 16 fig. 5 (d genita- lia); Szöcs, 1965: 78. Stigmella spinosella; Klimesch, 1951: 62; Gerasimov, 1952: 260; Hering, 1957: 835, fig. 518 (mine); Kli- mesch, 1961: 759; Lhomme, 1963: 1194; Emmet, 1970b: 121, 122, fig. 1. Dechtiria spinosella; Emmet, 1971: 244. Trifurcula (Ectoedemia) spinosella; Johansson, 1971: 245, Ectoedemia spinosella; Bradley et al., 1972: 2; Emmet, 1974a: 79, 80; Borkowski, 1975: 493; Emmet, 1976: 192, pl. 6 fig. 8, pl. 12 fig. 23; van Nieuker- ken, 1982: 108, fig. 8 (mine). Diagnosis: E. spinosella is externally similar to the angulifasciella complex, but is smaller, has a fuscous collar and the male has a brown hair-pencil surrounded by some brown lamellar scales. The female can be separated from atricol- lis by shorter signa with smoother, more uni- formly curved outline. See for separation from mahalebella under that species. Description. Male (fig. 84). Forewing length 1.44—2.20 mm (1.87 + 0.15, 29), wingspan 3.2—4.9 mm. Head: frontal tuft orange to orange fuscous, sometimes completely fuscous; collar fuscous. Antenna with 24—30 segments (26.8 + 1.7, 18). Thorax and forewings blackish fuscous with medial silvery fascia, slightly concave at inner margin. Hindwing with brown hair-pencil, sur- rounded by a small patch of brown, lamellar scales. Underside of forewing with a tuft of long grey or white hairscales, arising near costal reti- naculum. Female. Forewing length 1.52—2.24 mm (1.85 + 0.16, 34), wingspan 3.4—5.0. Antennae with 21—26 segments (22.5 + 1.1, 29). Hind- 76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 wing without brown patch, forewing without tuft. Male genitalia (figs. 137, 138, 277, 325, 395). Capsule length 193—219 um (207.9 + 8.9, 8). Tegumen produced into broad and truncate pseuduncus. Gnathos (fig. 325) divided, with short, rounded distal element, and basal part with serrate margin. Valva (fig. 277) length 133—150 um (142.5 + 5.5, 8), inner margin slightly sinuous to almost straight, tip pointed. Aedeagus (fig. 395) 231—253 um (242.1 + 7.6, 8), with single, or bifid, pointed carinae. Female genitalia (figs. 217, 218, 468). T7 without a row of setae. T8 with two lateral patches of scales and several setae (at least 4). Anal papillae with 6—11 setae. Vestibulum with incomplete vaginal sclerite, a spiculate pouch with indistinct spines. Corpus bursae 440—550 um, completely covered with small pectinations or minute spines; signa slightly dissimilar, ovoid, with smooth, uniformly curved outline, longest 249—373 um (312.4 + 44.6, 9) shortest 227—330 um (283.3 35.4, 9), 2.4—3.5 X as long as wide. Ductus spermathecae with 2—2” convolutions. Larva. Greenish white, with distinct brown ganglia. Head light brown. Ventral plates ab- sent. Biology. Hostplants. Prunus spp., in central and north- ern Europe only on P. spinosa L., in the south also recorded from P. domestica L., P. cerasifera Ehrh., P. fruticosa Pallas (to be confirmed), P. dulcis (Miller) (Greece). Mine (figs. 508, 509). Egg on leaf underside, close to mid-rib, or less often lateral vein; occa- sionally on leaf-margin. Early mine much con- torted narrow gallery, filled with reddish frass, later becoming elongate blotch with dispersed black frass, often very compact. Life history. Univoltine. Larvae from end of July to October, most abundant in September, but in southern Greece some mines were va- cated already by mid June. Adults in June and July (occasionally May). Distribution (fig. 536). Widespread in central Europe, but more lo- calised northwards, occurring mainly on sun- exposed hills, or near coast (in England). Proba- bly widespread in mediterranean area, but not yet recorded from Iberian Peninsula, the medi- terranean islands, and most of the Balkan. Borkowski (1975) did not mention E. spinosella from Poland, but the specimens cited below, collected by Hering, indicate its presence in Po- land. Recently Buszko (in litt.) found it also in Poland. Remarks. Before De Joannis discovered this species in France, it had been mistaken several times for E. atricollis. Named as such, specimens which were collected by Eppelsheim in Pfalz can be found in many collections. To my knowledge E. atricollis has never been found on Prunus spino- sa. Greek specimens reared from Prunus dulcis (on which itis locally almost a pest) differ slightly in head-colour and female signa (usually shorter), but electrophoretically they appeared to be indistinguishable from normal spinosella (Menken, in preparation). Material examined: 52 6, 59 ©. — Austria: 1 6, Dürnstein, e.l. 2.vi. 1936, J. Klimesch; 1 2, Kloster- neuburg, Freiberg, e.l. 15.vi.1941, Preissecker; 2 à, 1 9, Mödling, e.l. 7—25.v.1938, Preissecker; 1 4, Neu- Aigen, Schmidawiesen, e.l. 18.v.1937, Preissecker (NMW). — France: 3 d, 4 2 (lecto- and paralecto- types), Vannes, 18.vi, 2.vu, prunetier (= Prunus spin- osa), Joannis (MNHN). — Germany, West: 2 9, Grünstadt, Pfalz, Eppelsheim (MHUB, NMW); 3 9, Pfalz, e.l. 1893, 1894, Prunus spinosa, Eppelsheim (MHUB, NMW); 3 6, no data (probably Pfalz, Eppelsheim) (ZMA); 1 d, 2 ©, Lemberg-Zufth., Württemberg, e.l. 5—10.v.1939, Prunus spinosa, A. Worz (LNK, coll. Johansson). — Great Britain: 1 9, Puddle Dock (Essex), e.l. 4.v1.1982, Prunus spinosa, A. M. Emmet (ZMA). — Greece: 31 6, 25 ©, Arák- hova (Voiotia), 950 m, el. 2.v.—8.v1.1981, Prunus dulcis, 27—29.ix.1980, S. B. J. Menken, E. J. van Nieukerken (ZMA, ZSMK). — Hungary: 1 d, Badac- sony, e.l. 10.v1.1969, Prunus spinosa, J. Szöcs; 1 6, 3 ©, Torokbalint (W. of Budapest), el. 20.vi— 16.vu.1955, Prunus spinosa, J. Sz6cs (TMAB). — Netherlands: 3 d, 5 2, Gulpen, el. 7—16.v1.1980, Prunus spinosa, E. J. van Nieukerken; 1 5,1 9, 2 km NE Wijlre, Vrakelberg, e.l. 11—13.vi. 1980, Prunus spinosa, E. J. van Nieukerken; 3 d,9 2, Woods W. of Wijlre el. 22.vi—6.v11.1982, Prunus spinosa, Alders, Van Nieukerken (ZMA). — Poland: 1 d,2 ©, Krosna Odr. (Crossen a. Oder), el. 7—12.v1.1930, Prunus spinosa, Hering (MHUB). Mines. — On Prunus spinosa. — Austria: Gumpoldskirchen. — France: Villefranche-de-Con- tlent. — Germany, West: Kassel, 1.x.1946 (BMNH); Klotten. — Netherlands: Gulpen; Wijlre. — Poland: Bellinchen/Oder, W. of Chojna, 6.1x.1939, Hering; Krosna Odr. (Crossen/Oder), 1x.1929, Hering (BMNH). — Yugoslavia: Savudrija (Istria). On Pru- nus dulcis. — Greece: Arakhova; Dhelfoi; Kardhami- li. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 77 48. Ectoedemia (Ectoedemia) mahalebella (Klimesch, 1936) (figs. 85, 140, 141, 219, 278, 326, 396, 469, 510, 537) Nepticula mahalebella Klimesch, 1936: 207, 208, figs. 8, 9. Syntypes, Italy: Naturno, Vintschgau, e.l. 5—21.v. 1935, Prunus mabaleb, J. Klimesch (ZSMK) [not examined]. Nepticula mahalebella; Klimesch, 1940b: 190; Szöcs, 1965: 79. Stigmella mahalebella; Lhomme, 1945: 155; Kli- mesch, 1948: 72, figs. 47—49 (d genitalia); 1951: 62; 1961: 759; Lhomme, 1963: 1195. Nepticula (Dechtiria) mahalebella; Klimesch, 1950: 28, figs. 13—15 (mine, dé. genitalia, foodplant races). Stigmella (Dechtiria) mahalebella; Hering, 1957: 836 (mine). Ectoedemia mahalebella; Sz6cs, 1978: 266. Diagnosis: easily separated from related spin- osella by light collar, which is concolorous with frontal tuft (darker in spznosella), and absence of hair-pencil and special scales on hindwing of male, and by position and shape of signa in fe- male genitalia. Similar, but larger, angulifasciella separated by presence of hair-pencil and differ- ent shape of valva in male and signa in female. See also diagnosis of hexapetalae. Description. Male (fig. 85). Forewing length 1.92—2.40 mm (2.14 + 0.17, 7), wingspan 4.3—5.3 mm. Head: frontal tuft yellowish orange to ferrugi- nous; collar concolorous with or lighter than frontal tuft. Antennae with 26—32 segments (28.7 + 1.9, 7). Thorax and forewings blackish fuscous with medial silvery fascia, inner margin slightly concave. Hindwing without hair-pencil or costal bristles. Underside forewing with a tuft of long grey hair-scales, arising near costal retinaculum. Female. Forewing length 2.0—2.36 mm (2.15 + 0.10, 14), wingspan 4.4—5.2 mm. Antennae with 23—27 segments (24.5 + 1.2, 11). Male genitalia (figs. 140, 141, 278, 326, 396). Capsule length 201—214 um (4). Tegumen pro- duced into broad and truncate pseuduncus. Gnathos (fig. 326) divided into short, rounded distal part, and basal part with serrate margin. Valva (fig. 278) length 129—150 um (4), inner Margin straight, tip pointed; valva widest at base, constricted below tip. Aedeagus (fig. 396) 231—274 um (260.6 + 17.0, 5), with single or bifid, pointed carinae. Female genitalia (figs. 219, 220, 469). T7 without a row of setae. T8 with two lateral patches of scales and 6—8 setae. Anal papillae with 5—11 setae. Vestibulum with complete va- ginal sclerite, a spiculate pouch with indistinct spines. Corpus bursae long, 570—715 um, proximally covered with pectinations, distally with small spines; signa ovoid, almost similar, confined to proximal (posterior) half of corpus bursae, length 201—304 um (238.9 + 26.3, 19), 1.8—2.4 X as long as wide. Ductus spermathe- cae with 2'2—3 convolutions. Larva. Greenish white, with distinct brown ganglia. Head light brown. Ventral plates ab- sent. Biology. Hostplants. Prunus mahaleb L., on which most common, P. cocomilia Ten. (Greece), P. tenella Batsch (Hungary), P. fruticosa Pallas, P. avium L. and P. cerasus L. (Klimesch, 1950 and own data). Mine (fig. 510). Egg deposited on leaf-under- side, usually at or near margin, in French and Yugoslavian samples 99% at margin, but in Ital- ian and Greek samples up to 50% close to mid- rib or lateral vein. It is not yet clear if these min- es belong all to mahalebella. Early gallery nar- row, following leaf margin, or much contorted, filled with reddish frass; later abruptly changing into small roundish blotch, with blackish frass accumulated in centre. Life history. Univoltine. Larvae from late Ju- ly until mid-October. Adults in May and June (rearing data). Distribution (fig. 537). A southern European species, south and east of the Alps, including hot alpine valleys. Re- corded from Rumania as E. spinosella (Draghia, 1967). Remarks. The types have not been examined, but from Klimesch’s (1936) description, the identity of this species is clear. In central Europe E. mahalebella and spino- sella are clearly separated by their host-plants, but in the south, they could have overlapping hostplant ranges. More data are needed to con- firm this. Material examined: 12 6, 19 2. — Austria: 2 9, Bad Deutsch Altenburg, Pfaffenberg, el. 4 + 23.v1.1934, Weichsel (= P. mahaleb), Preissecker (NMW). — France: 1 6, 1 ©, St. Thibaud-de-Couz 78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 (Savoie), 500—700 m, e.l. 11—19.v1.1980, Prunus ma- haleb, E. J. van Nieukerken (ZMA). — Greece: 1 9, Parnassós Oros, NW Arakhova (Voiotía), plateau, 1150 m, el. 9—11.v.1981, Prunus cocomilia, S. B. J. Menken & E. J. van Nieukerken; 2 d 5 ©, Mt. Tim- fristös (Evritania) above Karpenission, 1200—1400 m, e.l. 21.v—25.vi.1981, Prunus cocomilia and P. maha- leb, S. B. J. Menken & E. J. van Nieukerken (ZMA). — Hungary: 3 ©, Budaörs, el. 3—7.vi.1971, Prunus mahaleb, J. Szöcs; 3 36, Budaörs, Csiki-hegyek, e.l. 20—27.v1.1962, Prunus mahaleb, J. Szöcs; 1 2, Bu- daòrs, Uthegy, e.l. 14.v1.1973, Prunus tenella, J. Szócs (TMAB). — Italy: 1 6, 1 ©, Trento, Goccladoro, e.l. iv.1946, Prunus mahaleb, J. Klimesch (ZMA). — Yu- coslavias 5 @ 5 2, Sele, 4 lam Ss Crikvenica (Biagio), acd, mimes W558, el Hi 3.v.1984, Prunus mahaleb, J. J. Boomsma & E. J. van Nieukerken (ZMA). Mines. — On Prunus avinm. — Italy: Guilliana near Savona, 17.1x.1944, J. Klimesch (BMNH); Fras- cati, 17.x11.1941, Groschke (BMNH). On Prunus co- comilia. — Greece: Oiti Oros, SW Ipáu (Fthióus); Oiti Oros, NE Strémi (Fokis); Mt. Timfristós above Karpenision; Parnassos Oros, NW Arakhova (Voio- tia). On P. fruticosa. — Austria: Hundsheimer Berg near Hainburg. On P. mahaleb — Austria: Hainburg- er Berge (BMNH). — France: St. Thibaud-de-Couz (Savoie); Modane (Savoie); Villefranche-de-Conflent (Pyr. Or.). — Greece: Mt. Timfristos above Karpen- ision; Kastraki (Trikala). — Italy: Avezzano (Lazio); Brenzone, x.1943, Groschke (BMNH); Susa, Pie- monte, 20.viii.1960, 1.1x.1964, Jackh (BMNH). — Yugoslavia: Crikvenica (Croatia); Novi Vinodolski (Croatia). The Ectoedemia occultella group This group comprises two closely related spe- cies, mining in Betulaceae. They differ from all other described Ectoedemia s.str. species by the absence of a cilia-line and the concolorous black cilia. See further the descriptions. The larvae are yellow and possess ventral plates. This group occurs also in North America (E. lindquisti (Freeman)) and in Japan. 49. Ectoedemia (Ectoedemia) occultella (Linnaeus, 1767) (figs. 4, 5, 86, 87, 139, 221, 279, 397, 405, 470, HU, SUZ, 550) Phalaena (Tinea) occultella Linnaeus, 1767: 899. Syn- types, Sweden: Hammerby, Linnaeus (depository unknown, probably lost) [not examined]. Tinea strigilella Thunberg, 1794: 87. Lectotype © (designated by Robinson & Nielsen, 1983), Swe- den: [Uppsala], Gedner, Genitalia slide RJ 751A (Zoological Institute, Uppsala) [not examined] [Synonymised by Robinson & Nielsen, 1983]. ? Tinea mucidella Hubner, [1814—1817]: pl. 65 fig. 435. Syntypes, [Europe] (depository unknown [not examined]. Tinea mediofasciella Haworth, 1828: 584. Lectotype 3 (here designated), [England: London], ex Ha- worth coll., Stainton coll., Genitalia slide 22608 (BMNH) [examined]. Syn. nov. Lyonetia argentipedella Zeller, 1839: 215. Lectotype 2 (here designated) [Poland: Glogow (Glogau)], 28.v.[18]35, Zeller, Walsingham coll. 1910—427, 101267, Genitalia slide 22600 (BMNH) [exam- ined]. [Synonymised by Robinson & Nielsen, 1983]. Lyonetia argentipedella; Tengström, 1848: 152. Nepticula argentipedella; Heyden, 1843: 208; Zeller, 1848: 316, 317; Stainton, 1849: 29; 1854: 303; Herrich-Schäffer, 1855: 353; Frey, 1856: 386, 387; 1857: 421, 422; Stainton, 1859: 435; 1862: 212—219, pl. 10 fig. 2; Heinemann, 1871: 218; Nolcken, 1871: 780; Wocke, 1871: 338; 1874: 101; Heinemann & Wocke, 1877: 754, 755; Snel- len, 1882: 996, 997; Sorhagen 1886: 307; Meyrick, . 1895: 721; Tutt, 1899: 289—291; Rebel, 1901: 225; Meess, 1910: 478, pl. 91 fig. 66; Sorhagen, 1922: 48, pl. 2 fig. 46; Meyrick, 1928: 858; Peter- sen, 1930: 66, fig. 82 (d genitalia); Szócs, 1965: 64. [no genus] argentipedella; Herrich-Schaffer, [1853]: pl. 105, fig. 834. Dechtiria argentipedella; Beirne, 1945: 205, fig. 62 (d genitalia). Stigmella argentipedella; Klimesch, 1951: 61; Gerasi- mov, 1952: 226; Klimesch, 1961: 758; Lhomme, 1963: 1188. Stigmella (Dechtiria) argentipedella; Hering, 1957: 179, fig. 124 (mine). Nepticula (Dechtiria) argentipedella; Szöcs, 1968: 226 (biology). Trifurcula (Ectoedemia) argentipedella; Johansson, 1971: 245. Ectoedemia argentipedella; Bradley et al., 1972: 2; Borkowski, 1975: 493; Emmet, 1976: 197, pl. 6 fig. 9, pl. 12 fig. 28; van Frankenhuyzen & de Vries, 1979: 129—135, figs. (biology). Trifurcula argentipedella; Karsholt & Nielsen, 1976: 18. Microsetia mediofasciella; Stephens, 1829: 208; 1834: 268. ? Elachista mucidella; Treitschke, 1833: 179. Ectoedemia occultella; Robinson & Nielsen, 1983: 221,222. Diagnosis: easily distinguished from other Ectoedemia spp. (except minimella), by com- pletely jet-black colour of thorax and forew- ings, (except fascia), including cilia, and absence of cilia-line. Separated from Stigmella species by medial fascia (usually postmedial in Stigmella) and collar, consisting of hair-scales, instead of lamellar scales as in Stigmella. Separated from very similar minimella by presence of group of Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 79 white scales on underside of forewing in male, and by light coloured head in female. See also minimella. Description. Male (fig. 86). Forewing length 2.36—3.44 mm (2.85 + 0.33, 23), wingspan 5.1—7.5 mm. Head: frontal tuft black, often mixed with some fuscous or ochreous scales; collar black. Anten- nae with 31—42 segments (35.6 + 2.9, 19). Thorax and forewings completely jet-black, less coarsely scaled than in other Ectoedemia spe- cies, with a rather broad, almost straight dull white fascia, sometimes slightly constricted in middle. Hindwing with a relatively long white hair-pencil. Underside forewing with a small elongate patch along costa with narrow white scales, often difficult to see. Female (fig. 87). Forewing length 2.56—3.84 mm (3.28 + 0.39, 20), wingspan 5.7—8.4 mm. Head: frontal tuft yellowish to yellowish Orange, sometimes mixed fuscous; collar yel- low. Antennae with 27—32 segments (29.4 + 1.6). Patch of white scales on underside fore- wing absent. Male genitalia (figs. 139, 279, 397, 405). Cap- sule length 313—390 um (353.6 + 27.4, 10), very large comparing with other Ectoedemia (s.s.) species. Tegumen produced into long ta- pering, pointed pseuduncus. Gnathos (fig. 327) with relatively broad, blunt central element. Valva (fig. 279) length 236—279 um (245 + 8.3, 6), outer margin strongly convex, inner margin slightly concave, almost straight; tip pointed, pointing posteriorly. Aedeagus (figs. 397, 405) 304—351 um (326.8 + 17.5, 12), carinae each divided into several blunt ending digitate pro- cesses, number variable; vesica with many small, triangular cornuti only. Female genitalia (figs. 221, 470). T7 without row of setae. T8 with two groups of scales and 3—5 setae. Anal papillae confluent, in total with 18 to 40 setae. Vestibulum with vaginal sclerite, and a dorsal spiculate pouch with very few min- ute spines only. Corpus bursae 495—580 um, with pectinations closely set in two lateral bands, at some distance from signa; signa dissi- milar, one reaching vestibulum, longest 214— 334 um (275.3 + 34.1, 9), shortest 180—266 um (221.0 + 27.9, 9), 2.2—3.3 X as long as wide. Ductus spermathecae with 2/3 convolu- tions. Larva. Pale yellowish white, ganglia not very conspicuous. Head light brown. Penultimate stages with 12 black ventral plates. Biology. Hostplants. Betula spp. Occurring on al na- tive Betula spp. in Europe and many species in botanical gardens (Buhr, 1935, Van Franken- huyzen & De Vries, 1979). In northern Finland it has been found mining both on Salix pentan- dra L. and Betula, but no adults have yet been reared from Salix (J. Kyrki, pers. comm.). Mine (figs. 511, 512). Egg on leaf underside, rarely on upperside. Mine large blotch, often al- most circular, with black circular blotch in mid- dle, caused by staining of both epidermis layers; frass black, irregular, but usually accumulated under and near blotch. Mine does not start as gallery, young mines consist of black blotch only, through which larva cannot be seen. Life history. Univoltine. Larvae feed slowly during long period, from the end of June to ear- ly November. Complete mines with mature lar- vae can occasionally be found from late July to August, but are most common in September and October. Adults fly in May and June. See de- tailed description by Van Frankenhuyzen & De Vries (1979). Distribution (fig. 530). One of the commonest and most widespread Ectoedemia species in Europe, occurs in almost all places where birch grows. In southern Eu- rope probably in mountains only, and recorded from Etna, Sicily. Remarks. This species has long been known as E. ar- gentipedella (Zeller), but Robinson & Nielsen (1983) showed that this is a junior synonym of occultella Linnaeus. The type series of Tinea mediofasciella Ha- worth comprises five specimens, representing several species, including Bucculatrix, Stigmella, and one Ectoedemia. E. mediofasciella was pre- viously incorrectly synonymised with woolho- piella Stainton (= minimella), probably on the basis of the single Ectoedemia specimen, se- lected here as lectotype. Examination of the genitalia, which had not earlier been dissected, however, showed it to be occultella. The identity of Tinea mucidella Hübner is still unknown, this synonymy has been sug- gested by Zeller (1839) in his description of ar- gentipedella. E. lindquisti (Freeman, 1962), described also by Wilkinson & Scoble (1979) and Wilkinson & Newton (1981) is extremely similar to occultella in the adult and larval stage and in its life history 80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 (Lindquist, 1962). The only difference seems to be the absence of a patch of white scales on the underside of the forewing of the males. The al- lozyme differences are also small (Menken, in preparation), so it is probable that lindquisti and occultella are vicariant forms, and hence differ- ent subspecies. Material examined: 102 d, 88 9, 141 ex. — Aus- tria: 1 ©, no further data (RMNH). — France: 3 d, 19, Pralognan (Savoie), 1450 m, e.l. 21—23.v.1980, E. J. van Nieukerken (ZMA). — Germany, West: 1 d, 1 2, Alendorf, 8 km S. of Blankenheim (N.-Westf.), e.l. 10—18.v.1983, Alders & Van Nieukerken (ZMA); 1 ©, Stuttgart (MHUB). — Germany, East: 1 ©, Berlin, Bot. Garten, 12.v.1947, Hering; 2 6, 1 ©, Berlin Fin- kenkrug, 1918—1932, Hering; 1 2, Berlin Frohnau, 18.11.1920, Hering; 3 5, 6 2, Potsdam, 13.1v.1886, 15—20v.1898, Hinneberg; 1 ex., Rachlau, 1884, Schutze (MHUB). — Great Britain: 1 4,5 ©, Brom- ley (London), 7—11.v.1939, S. Jacobs (ZMA); 1 d (lectotype of mediofasciella, see above). — Nether- lands: 82 d, 61 ©, 140 ex. from following localities: Arnhem; Epen; Geulhem; ’s-Gravenhage; Hilver- sum; Hoge Veluwe; Kerkrade; Kortenhoef; Korten- hoefse Plassen; Kosberg; Loosduinen; Meijnweg; Neerbosch; Nunspeet; Rockanje; Schin op Geul; Schinveld; Slenaken; Wageningen; Wijlre; Winters- wijk; Zwanewater (RMNH, ZMA, AFW, coll. Huis- man, coll. Kuchlein). — Poland: 1 © (lectotype of ar- gentipedella, see above); 2 ©, Obernigk, 11.1869; 1 à, 2 ©, Wroclaw (Breslau), 11.1869, Wocke (MHUB). — Switzerland: 1 6, 2 2, Zürich (MHUB). — No Lo- cality Data: 5 d,2 © (ZMA, RMNH, MHUB). Material of lindquisti examined. — Canada: 6 d, 3 9, Ontario: Awenda Prov. Park, Penetang, Simcoe Co., mines 24.viii.1981, Betula pe Evans, e.l. 2—8.v1.1982 (ZMA); USA: 2 6, 1 2, Maine, Bethel, 29.v1.1946, A. F. Braun (USNM). Mines. — Austria: Gramatneusiedl; Hermagor; Mühlleiten; Lavamünd. — Belgium: Bolderberg, Zolder. — France: Le Hohwald; Pralognan. — Ger- many, West: Alendorf; Oberstadtfeld. — Great Brit- ain: Brentwood; Grays; New Forest. — Hungary: Budapest. — Italy: Naturno; Tolmezzo; Trento. — Netherlands: many localities. — Yugoslavia: Fuzine, SW Delnice. 50. Ectoedemia (Ectoedemia) minimella (Zetterstedt, 1839) comb. n. (figs. 88, 142, 222, 280, 328, 398, 406, 414, 415, 513, 531) Elachista minimella Zetterstedt, 1839: type ® (here designated), Norway: Nordland, Bjorkvik, 14.vii, Zetterstedt, Genitalia slide RJ (Zoological Institute, Lund, Sweden) [examined by R. Johansson]. Nepticula woolhopiella Stainton, 1887: 262. Lecto- type 2 (here designated), Great Britain: Tarring- 1011. Lecto- ton, 29.vi.1887, e.l. birch, Wood, Genitalia slide 11362 (BMNH) [examined]. Syn. nov. Nepticula viridicola Weber, 1937: 211, 212, fig. 1. Lectotype 6 (here designated), Switzerland: Sim- plon, 1970 m, mines 19.1x.1936, Alnus virid., Z. 2606, Weber, Genitalia slide ETH 1241 (ETHZ) [examined]. Syn. nov. Nepticula argentipedella [partim]; Meyrick, 721. Nepticula woolhopiella; Tutt, 1899: 292, 293; Rebel, 1901: 225; Meess, 1910: 478; Meyrick, 1928: 858; Petersen, 1930: 67. Dechtiria woolhopiella; Beirne, 1945: 205, fig. 63 (d genitalia). Stigmella woolhopiella; Gerasimov, 1952: 270; Klı- mesch, 1961: 758; Borkowski, 1969: 100. Stigmella (Dechtiria) woolhopiella; Hering, 1957: 181, fig. 118b (mine). Trifurcula (Ectoedemia) woolhopiella; Johansson, 1971: 245. Ectoedemia woolhopiella; Borkowski, 1975: 493. 1895: Stigmella viridicola; Klimesch, 1948: 70, figs. 43, 44 (8 genitalia); 1951: 61; Hering, 1957: 66, fig. 37a (mine); Klimesch, 1961: 758. Ectoedemia woolhopiella viridicola; Borkowski, 1975: 494, [Ectoedemia mediofasciella; Bradley et al., 1972: 2; Emmet, 1973: 282, 283; 1976: 197, pl. 6 fig. 12; pl. 12 fig. 29; Van Nieukerken, 1982: 107, 108, fig. 7 (mine). misidentification]. [Trifurcula mediofasciella; Karsholt & Nielsen, 1976: 18. misidentification]. Diagnosis: extremely similar to occultella, for external differences see under that species. Male genitalia can be separated by smaller size, pres- ence of large elongate cornuti and shape of gna- thos. Female genitalia extremely difficult to sep- arate, but minimella has usually shorter and wider signa, although there is some overlap. Description. Male (fig. 88). Forewing length 2.32—2.72 mm (2.54 + 0.11, 14), wingspan 5.1—6.1 mm. Head: frontal tuft black; collar black. Antennae with 35—42 segments (37.5 + 2.2, 14). Thorax and forewings completely jet-black, less coarse- ly scaled than in other Ectoedemia species, with a rather broad, almost straight, dull white fascia, sometimes slightly constricted in middle. Hindwing with a greyish hair-pencil, slightly shorter than occultella. Underside of forewing without white scale patch. Female. Forewing length 2.28—3.04 mm (2710 = 10:24. 16), wingspan) 55|——6-Geamame Head: frontal tuft black, or mixed with yellow and fuscous scales, sometimes completely yel- low on frons, but always black on vertex; collar Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 81 black. Antennae with 23—29 segments (26.3 + 1.9, 14). Male genitalia (figs. 142, 280, 328, 398, 406, 414, 415). Capsule length 296—321 um (307.9 + 11.6, 6). Tegumen (figs. 414, 415) produced into long tapering, pointed pseuduncus. Gnathos (fig. 328) with narrow, truncate central element. Valva (fig. 280) length 214—227 um (221.0 + 4.2, 7), outer margin strongly convex, inner margin slightly concave, almost straight; tip pointed, pointing posteriorly. ons (figs. 398, 406) 283—309 um (297.1 + 10.0, 6), carinae each divided into several blunt ending processes, number variable; vesica with about 20—22 long, needle shaped cornuti at right side, and many smaller cornuti in remaining part of vesica. Female genitalia (figs. 222, 471). T7 without row of setae. T8 with two groups of scales and about 4 setae. Anal papillae confluent, with 23—32 setae in total. Vestibulum with vaginal sclerite, and dorsal spiculate pouch with very few minute spines only. Corpus bursae 440— 550 um, with pectinations in two lateral bands, at some distance from signa; signa dissimilar, longest 176—279 um (240 + 37.4, 7), shortest 167—231 um (199.6 + 24.9, 7), 2.0—2.4 X as long as wide. Ductus spermathecae with 2'2—3 convolutions. Larva. Pale yellow to yellowish white, with distinct brown ganglia. Head light brown. Pen- ultimate instars with 12 black ventral plates. Biology. Hostplants. Betula spp., usually on B. pubes- cens Ehrh., or in Scandinavia B. nana L., less common on B. pendula Roth. In the Alps com- mon on Alnus viridis (Chaix) DC. in Lam. & DC., which it seems to prefer even in the pre- sense of Betula. In the west of Great Britain also recorded from Corylus avellana L. Mine (fig. 513). Egg deposited on leaf-under- side. Early mine much contorted gallery, with dispersed frass, staining leaf brown; later abruptly enlarges into elongate blotch, which often fills the space between two veins; dispers- ed black frass. Life history. Univoltine. Larvae found from July to September, occasionally October in south, but most abundant in August and early September. Adults fly in May and June. Distribution (fig. 531). Common and widespread in Scandinavia and locally in the Alps, but elsewhere very local and always less common than occultella. Not yet re- corded from Belgium, Spain (to be expected in Pyrenees) or Yugoslavia (Alps). Remarks. This species has been known since 1972 (Bradley et al.) under the name E. mediofasciel- la, but this was apparently based on a misinter- pretation of the type, which in fact belongs to occultella. The first available name appears now to be minimella Zetterstedt, a name of which the identity was hitherto unknown. The two speci- mens mentioned by Zetterstedt (1839) are both in Lund, and were examined by R. Johansson, who kindly communicated us his observations. The specimen labelled “minimella 2” is identi- cal with woolhopiella, and selected as lectotype. The other specimen, described as variety, is a fe- male of E. (Fomoria) weaveri (Stainton). There seem to be no grounds for regarding viridicola as a subspecies (Borkowski, 1975), since it is not geographically or morphologically separate, and shows no differences in allozyme pattern (Menken, in preparation). The fact that it often feeds only on Alnus, even in the pres- ence of Betula might be explained by the ovipo- sition preference of females, which they cannot follow in other parts of its range where Alnus viridis is absent. It would be interesting to study minimella populations — if present — in north- ern Siberia or Corsica, where other subspecies of A. viridis occur. The character given by Beirne (1945) to sepa- rate minimella from occultella is incorrect and probably based on an artifact. It is questionable if the genitalia, depicted by him, belong to muni- mella, since he did not figure the characteristic cornuti. Material examined: 17 d, 22 2. — Austria: 2 9, Gr. Glockner, Guttal, 2000 m, e.l. iv.1944, Alnus viri- dis, J. Klimesch (ZMA). — France: 1 d, 1 ©, Pralog- nan (Savoie), 1450 m, e.l. 17—19.v.1980, Alnus viri- dis, E. J. van Nieukerken (ZMA). — Great Britain: 1 9 (lectotype, see above). — Italy: 1 ©, Riva di Tures (Rain in Taufers), Knuttental, 1800 m, 14.vi.1976, G. Derra (coll. Derra). — Netherlands: 2 2, Lochem, Ampsensche Veld, e.l. 16—18.iv.1983, Betula pubes- cens, E. J. van Nieukerken; 1 2, Rockanje: Voornes Duin, e.l. 20—21.v.1980, Betula pubescens, E. J. van Nieukerken (ZMA). — Norway: 1 6, 1 ©, Alta (Al- ten), 1.vu, Staudinger (MHUB); 4 d, 3 ©, Grovuda- len, 900 m, 62.27 N, 8.54 E, el. 5—22.v.1981, Betula pubescens, E. J. van Nieukerken; 1 6, 1 2, 2 km E. Oppdal, 650 m, e.l. 7—8.v.1981, Betula pubescens, E. J. van Nieukerken; 6 6,6 2, 11 km W. Rennebu, 600 82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 m, el. 30.iv—12.v.1981, Betula pubescens, E. J. van Nieukerken (ZMA). — Switzerland: 1 d, 1 © (lecto- and paralectotype of viridicola), Simplon, 1970 m, mine 19.1x.1936, Alnus virid., Weber (ETHZ); No lo- cality data: 3 d, 2 2, bred in captivity (from Norwe- gian material), el. 8—21.v.1982, Betula (ZMA). Mines. — On Alnus viridis. — Austria: Lavamund. — France: Pralognan. — Italy: Trento. — Switzer- land: near Genève. On Betula nana. — Norway: Grevudalen. On Betula pendula. — Germany, West: Oberstadtfeld. On Betula pubescens. — France: Pra- lognan. — Netherlands: Dalfsen; Griendtsveen; Den Ham; Lochem; Mariénberg; Oostvoorne; Ootmar- ‘sum; Rockanje; Vilsteren; Vorden. — Norway: Grovdal; Grovudalen; Hoem; Oppdal; Rennebu. NAMES OF DOUBTFUL STATUS, PROBABLY BELONGING TO ECTOEDEMIA Nepticula bistrimaculella Heyden, 1861: 40. According to Dr. H. Schröder (in litt.) there is no type-material of this species left in the Heyden collection in Frankfurt. From the de- scription it seems to belong to the subbimaculel- la complex and to feed on Betula. Most likely this refers to an unusual case of xenophagy of either ering: or subbimaculella. Nepticula gilvella Rossler, 1866: 395; 1881: 338. No material of this species is present in the Rossler collection in Wiesbaden (Dr. M. Geis- thardt, in litt.) nor in Strasbourg (Dr. J. Matter, in litt.). The description is vague, so the identity of this species-remains obscure. It could belong to one of the Quercus feeding Ectoedemia spe- cies. CATALOGUE OF HOSTPLANTS OF WESTERN PALAEARCTIC ECTOEDEMIA (Occasional occurrence on unusual hostplants in brackets) SALICACEAE Salix fragilis L. E. intimella Salix caprea L. E. intimella Salix cinerea L. E. intimella Salix pentandra L. E. intimella, (occultella) Salix phylicifolia L. E. intimella Populus alba L. E. klimeschi, turbidella Populus canescens (Aiton) Sm. E. turbidella Populus tremula L. E. argyropeza Populus nigra L. E. hannoverella, (turbidella?) Populus X canadensis Moench. E. hannoverella BETULACEAE (incl. Corylaceae) Betula pendula Roth. E. occultella, minimella Betula pubescens Ehrh. E. occultella, minimella Betula nana L. E. occultella, minimella Alnus viridis (Chaix) DC. in Lam. & DC. Corylus avellana L. Carpinus betulus L. FAGACEAE Fagus sylvatica L. Castanea sativa Miller Quercus coccifera L. Quercus ilex L. and rotundifolia Lam. Quercus suber L. Quercus macrolepis Kotschy Quercus alnifolia Poech Quercus infectoria Olivier Quercus cerris L. E. minimella (E. minimella) E. (Zimmermannia) spec. E. liebwerdella E. (Zimmermannia) spec., E. albifasciella, heringi E. cf. algeriensis, haraldı, suberis, andalusiae, cf. caradjai E. (Zimmermannia) spec., E. algeriensis, ilicis, heringella, baraldi, suberis E. haraldi, ilicis, suberis E. aegilopidella E. heringella, alnifoliae E. cf. caradjai ? E. caradjai, gilvipennella, cerris, (subbimaculella), liechtensteini, phyllotomella Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia Quercus petraea L. s.l. Quercus robur L. Quercus frainetto Ten. Quercus pyrenaica Willd. Quercus pubescens Willd. s.l. Quercus faginea Lam. Quercus ebrenbergi Kotschy ULMACEAE Ulmus spp. ROSACEAE Spiraea media Franz Schmidt Filipendula vulgaris Moench Agrimonia eupatoria L. Aremonia agrimonioides (L.)DC. Rubus chamaemorus L. Rubus arcticus L. Rubus saxatilis L. Rubus caesius L. Rubus fruticosus L. aggr. Rubus ulmifolius Schott Rosa spp. Sanguisorba officinalis L. Sanguisorba minor Scop. Potentilla erecta (L.) Rauschel Potentilla sterilis (L.) Garcke Fragaria vesca L. Fragaria moschata Duchesne Pyrus communis L. Malus sylvestris Miller ? Sorbus sp. Mespilus germanica L. Crataegus laevigata (Poiret) DC. Crataegus monogyna Jacq. Prunus dulcis (Miller) Prunus tenella Batsch Prunus cerasifera Ehrh. Prunus spinosa L. Prunus domestica L. Prunus fruticosa Pallas Prunus cocomilia Ten. — Prunus avium L. _ Prunus cerasus Prunus mahaleb ANACARDIACEAE Pistacia terebinthus L. STAPHYLEACEAE Staphylea pinnata L. E. caradjai, quinquella, nigrosparsella, albifasciella, subbimaculella, heringi E. atrifrontella, longicaudella, quinquella, albifasciella, (contorta), subbimaculella, heringi E. caradjai, E. albifasciella complex E. subbimaculella E. atrifrontella, caradjai, nigrosparsella, pubescivora, contorta, subbimaculella, heringi, (? liechtensteini) ? E. atrifrontella, E. cf. suberis, heringi E. caradjai E. amani, preisseckeri E. spiraeae E. hexapetalae, angulifasciella E. agrimoniae, (? rubivora, ? arcuatella) . agrimoniae . rubivora rubivora rubivora rubivora . rubivora, erythrogenella erythrogenella . angulifasciella . angulifasciella . angulifasciella . arcuatella arcuatella arcuatella arcuatella atricollis atricollis atricollis „atricollis atricollis atricollis spinosella mahalebella spinosella ?, (atricollis) spinosella spinosella, (? mahalebella, ? atricollis) mahalebella .mabalebella .atricollis, mahalebella .mahalebella . mahalebella, (atricollis) tay Ors try Es Ens Ees Pr Ors Ems Ory Ems Eri oo em EM E. terebinthivora E. atricollis 83 84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 È N° à 2 N e NG OP RO I LE LAS ERS STILO EN SME x ss I x © Ke ; NN & Ss © Le 4 AS È 33* 26 Bas 36* 23A Ja = 34L 38 28 37* 25 24 23 11 10 9%* 8 ] 20018 > 21 k 20 3 2 1 Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 85 PHYLOGENY I have attempted to reconstruct the phyloge- ny of Ectoedemia, using the cladistic approach as outlined by Hennig (1966) and refined amongst others by Wiley (1981). Many difficulties arose with the assessment of the polarity of character states, especially within the subgenus Ectoedemia, since at first sight there seemed to be no correlation at all between the character distributions. This means that there is a considerable amount of either homo- plasy, secondary reduction, reversal or cases of underlying synapomorphies (Saether, 1979), which require many ad hoc statements to ex- plain apparently conflicting evidence. It must be stressed that many of such characters are rela- tively simple morphological structures, which therefore might have a simple genetic basis. If so, reversals and parallelisms could appear quite often in the course of evolution. Therefore it is not always feasible to use parsimony, where on- ly the number of “ad hoc” statements counts, not their quality. The most parsimonious clado- gram should only been chosen after some qual- ification or weighing of the “ad hoc” statements. For instance reduction of a simple structure is a much more likely event than the parallel development of a complex structure. Another weak point in the phylogeny pre- sented and discussed below, is that several mo- nophyletic groups are defined by one character only. Moreover these characters are frequently suspected to be homoplasies, but similarity both in morphology and biology often coincides with the groups defined in the cladogram, and al- though part of this similarity might be based on plesiomorphies, it is also very likely that apo- morphies, which can at present not be defined easily, play an important role in this similarity. An extension of this analysis with Nearctic and eastern Palaearctic species, and larvae could res- olve some of the existing uncertainties. The following analysis has been carried out by hand and as a consequence of the high pro- KG NÉ Q v> NES SS EN Oe HECHT NO) at È GO KT LE ON à ES © os ES SS NS È SS LT OSE oo de Co x I SN S LELE ET Vela ELE gol 40* 11% 46 47 52 58 39 15% 5! 93 57 [AR 50 56 ~ 43 55% 42 6 * xD & 4 = sij 38% 49 * 35 3 34 L Figs. 1—3. Cladograms representing proposed phylogeny within Ectoedemia. Black squares denote apomor- phies; black dots characters with uncertain status. Character numbers explained in text; in-group parallelisms marked with an asterisk, frequent secondary loss denoted by L. Figs. 1 and 2 give two alternative phylogenies for the basic branching in the genus, fig. 3 details the right branch of figs. 1 and 2. 86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 portion of conflicting evidence, can only be re- garded as a very rough preliminary analysis, open to further tests. The supposed apomor- phies found with the outgroup-rule are given for pairs and groups of species. Autapomor- phies for single species are not given here. Cla- dograms representing the proposed phylogenies are presented in figs. 1—3. - Sistergroup and monophyly of Ectoedemia The sistergroup of Ectoedemia s.str. and E. (Zimmermannia) should be sought for amongst the taxa Fomoria Beirne, Laqueus Scoble or Etainia Beirne (van Nieukerken, in prepara- tion). For neither of them convincing arguments have been found, so for the following outgroup comparisons all these taxa together have been taken into consideration. The following apo- morphies support the monophyly of the two subgenera treated here and therefore corrobo- rate the earlier suggestion of monophyly based on character 1 only (Scoble, 1983). 1. Loss of uncus. — The classical character (Beirne, 1945; Scoble, 1983). The uncus is also absent in Holarctic species of Etainia, but since it is present in some South Afri- can Etainia species, it has probably been lost independently. An uncus is present in almost all other Nepticulidae. 2. Sensillum vesiculocladum blisterlike, not branched (van Nieukerken & Dop, in preparation). — A more or less similar sit- uation in some species of Fomoria is tenta- tively regarded as a parallelism. 3. Female with single sensillum vesiculocla- dum per flagellar segment (van Nieukerken & Dop, in preparation). — A unique char- acter, checked for many species, represent- ing all species groups. Subgenus Zimmermannia The Palaearctic and Nearctic species share a number of uniquely derived characters which demonstrate the monophyly of this subgenus, and therefore justify its re-establishment. 4. Larvae barkmining. — The basic feeding pattern in Nepticulidae larvae is leaf-min- ing. Although some other species make mines in bark of branches or shoots, only Zimmermannia larvae make mines in the bark of thick branches or trunks of trees, especially Fagaceae. This character’ led Hering (1940) to erect the genus Zimmer- mannia, but later authors doubted the va- lidity of this character to define a taxon (i.e. Wilkinson & Newton, 1981). In my opinion it is a sound autapomorphy for the subgenus. 5. Larval life lengthened, with 6—8 instars. — As a rule Nepticulidae larvae have four or five larval instars, with probably four as the most generalised condition (van Nieu- kerken & Jansen, in preparation). How- ever, this apomorphy is subject to some reservation as it is only known with cer- tainty for atrifrontella, liebwerdella and longicaudella. 6. Colour pattern of forewings largely lost, colour uniform or irrorate. — The pres- ence of light dots or fasciae is assumed to be the generalised condition in Nepticuli- dae. 7. Male hindwing with pronounced costal emargination. — The costal emargination, - unknown outside Zimmermannia, is asso- ciated with the relatively long hair-pencil. In some species with reduced or without hair-pencil, the emargination is absent. A hair-pencil is considered to belong to the ground-plan of Ectoedemia and 1s also pre- sent in several non-European species of Etainia, Laqueus and Fomoria. Therefore the reduction of the hair-pencil and hence of the emargination are thought to be secondary (character 18). 8. Large size of ventral carinae and corre- sponding dorsal fold of valva. — A pecu- liar feature, which is clearly seen in undis- sected genitalia. 9. Female with many long tactile setae on ter- gites 7 and 8. — This character needs in- vestigation in Nearctic species. It is proba- bly secondary reduced in amani and liguri- cella (character 19). 10. Bursa copulatrix extremely long and nar- row. 11. Margin of signa wider than individual cells. The available characters are insufficient to present a cladogram of the western Palaearctic species of Zimmermannia, however some sup- posed apomorphies for groups of species are given below and listed in table 1. 12. Aedeagus constricted. 13. Dorsal and dorsolateral carinae connected by rim. Characters 12 and 13 show the sister relationship between atrifrontella and liebwer- della. 14. Vesica with folded sclerotised plate. — This character is shared by the first three RT LE EEE TTT EE VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 87 Table 1. Data-matrix of some important characters in Ectoedemia (Zimmermannia) species. Species given with their number and first three letters of epitheton, characters and numbers refer to text. — 1 = character present (supposed apomorphy), 0 = character absent (either by plesiomorphy or secondary reduction), ? = status unknown. Sh ance hte Dar Me omsten brei DER heao Cha 2 3 4 5 6 7 8 12 1 1 (0) 0) (0) 0 0) 0) 13 1 1 (0) (0) (0) (0) 0 0 14 1 1 1 ? 0 (0) (0) 0 15 0 0 1 It 1 1 1 1 16 (0) 0 1 ? 1 1 1 1 1174 0 (0) (0) 0 1 1 0) (0) 18 0 (0) 0 (0) 0 0 1 il 19 (0) 0) 0) 0 (0) 1 0 1 species and therefore in conflict with the following characters. 15. Valva with inner (mesal) lobe. — This lobe is only slightly developed in longicaudella and nuristanica. 16. Ductus spermathecae with more than 312 convolutions. — The basic number is 2 to 3 convolutions, increase of this number oc- curred independently in various other groups. Characters 15 and 16 indicate a monophyly of the species 3 to 8, but they are both only slightly developed in longicaudella, which to- gether with character 14 make its position un- certain. The valval lobe is also only slightly de- veloped in nuristanica, but it is present in many Nearctic species, and it is therefore not at all un- likely that the lobe belongs to the ground-plan of Zimmermannia and has been lost in a few species. 17. Vesica with stout sclerotised cornutus. — Present in amani, monemvasiae and also in several Nearctic species, in which the cor- nutus bears also many secondary spines. It is not clear if this character is a homologue of the sclerotised plate (character 14) and hence part of the same transformation se- ries. In that case either 14 or 17 is invalid as autopomorphy. 18. Loss of hair-pencil (and costal emargina- tion) in 6. — See character 7. 19. Loss of long setae on abdominal tip in ©. — See character 9. Subgenus Ectoedemia s.str. The following characters of the female genita- lia are assumed to be apomorphic for the subge- nus: 20. Vestibulum with circular vaginal sclerite. — Vaginal sclerites are present in several other nepticulids and according to Scoble (1983) belong to the ground plan of Trifur- culini, but they usually have a different shape from the type here, which is unique for Ectoedemia s.str. It is only absent in spiraeae and agrimoniae. 21. Vestibulum with spiculate pouch. — Prob- ably correlated with 20, this is another unique character for the subgenus, which is absent in the same two species and hexape- talae, and less distinct or without spicules in some other species. 22. Vestibulum with patch of densely packed pectinations. — Shared by all species of the populella group, subbimaculella group, preisseckeri, terebinthivora and erythroge- nella. It is either another synapomorphy for the subgenus (fig. 1) or of a large part (fig. 2), but in both cases secondarily lost in many species. The subgenus also exhibits high uniformity in several other characters. For instance in the shape of the valva, the aedeagus and genital cap- sule; the general shape of the female genitalia and several biological characters. Yet it appears to be impossible to ascribe any of these similari- ties to straightforward apomorphies, indeed some of them are rather plesiomorphic. Some other features, which easily identify a species as belonging to Ectoedemia s.str. cannot be re- garded as belonging to the groundplan because they are absent in too many species, to explain them all as secondary losses. However, present evidence justifies the acceptance of Ectoedemia s.str. as amonophyletic entity. Subdivision of the subgenus into species groups is desirable, for coping with the large number of species. The aim has been to make monophyletic groups, but on the basis of the species treated here, it is difficult, and the subdi- vision only tentative. The groups used here are recognised by a combination of similarities in both morphological and biological characters. For some species which were hard to place, the biology provided the decisive factors, so that all groups recognised here feed on one hostplant family. Most of these groups are likely to be monophyletic, but at least one is suspected to be paraphyletic. With the characters given, I have presented two alternative phylogenies in figs. 1—3, but both still require many ad hoc 88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 statements. The characters used are discussed below and partly presented in the data-matrix in table 2. Characters are treated in the order in which they appear in the cladograms figs. 1 and 3, which I regard at present as the best alterna- tives. The sequence of species in the main body of the text also follows these cladograms. The populella group forms one of the best de- fined groups in Ectoedemia, with a high overall similarity and the following supposed apomor- phies: 23. Petiole or midrib miners. — Just as in the case of Zimmermannia, this feeding pat- tern is so unique and different from leaf- mining, that it can be safely regarded as an apomorphy for the populella group. With- in this group the petiole-mining is proba- bly more derived than the midrib mine of intimella in Salix, which could be the first step in the evolution from a “normal” leaf- mine into a petiole mine. Hence, the peti- ole-mine in Populus is regarded here as a further step in the transformation series and as such used with number 23A in fig. 1. A mine on Ostrya, strikingly similar to that of intimella, has been figured by Clemens (1872: figure on p. 27), but re- mains undescribed. 24. Hostplant: Salicaceae. — The character “hostplant” is difficult to interprete, but certainly useful in some cases. It is possible that oak (Quercus) is the ancestral host- plant for Ectoedemia s.str. because it is also the main hostplant for the sister-group, Zimmermannia. This explains the fact why two rather different groups mine in Quer- cus; they have retained their plesiomorphic hostplant. Salicaceae certainly seems to be a good apomorphy. In other leaf mining taxa, species feeding on Salicaceae are closely related (Stigmella, Phyllocnystis). 25. Denticles on spiculate pouch single, equal- ly spaced. — This character is diagnostic for the populella group, but it is impossible to decide if it is derived or ancestral. On grounds discussed above intimella is re- garded as the sister-species of the remaining Populus feeding species. The following charac- ter seems to be an apomorphy for turbidella and klimeschi. 26. Aedeagal carinae very well developed and large. — The total configuration of aedea- gus, and in fact the male genitalia as a whole is very similar in these two species. This character is of course inappropriate for the parthenogenetic argyropeza. On the basis of high similarity this species can be regarded as closely related to klimeschi, which might be the sexually reproducing ancestor of argyropeza. For the remaining species groups the follow- ing character is tentatively regarded as the only apomorphy: 27. Second and third larval instar with 12 ven- tral plates. — A unique character present in many species of Ectoedemia s.str., but again often absent from closely reihe spe- cies. It does not occur in the populella group and suberis group. It is supposed to be an apomorphy for the subgenus without the populella group in fig. 1 or for the sub- genus without the szberis group in fig. 2. It is either lost in the species in which it is ab- sent, Or it is an underlying apomorphy. E. preisseckeri has some affinities with the al- | bifasciella complex but since it lacks apomor- phies 28, 35 and 36 is placed here in a group of Its Own, as a sister-group of the remaining spe- cies. It is probably closely related to the Nearc- tic E. ulmella (Braun). All other species belong to one monophyletic entity on the basis of the following apomorphy: 28. Aedeagus with only one pair of carinae. — Within Ectoedemia s.l. the presence of sev- eral (2—4) pairs of carinae is widespread in the other subgenera, and therefore the ple- siomorphic condition on grounds of out- group argument. In cladogram fig. 1 the reduction to one pair is regarded as an apo- morphy for the remaining groups. The very similar configuration of the carinae in all species favours this solution, but a re- duction on several occasions cannot be ex- cluded and leads for instance to the clado- gram in fig. 2. In spiraeae the dorsal cari- nae are also lost. The species of the suberis group share the fol- lowing apomorphies: 29. Aedeagus very long in relation to capsule. 30. Signa oval. — The plesiomorphic condition of the signa seems to be narrow elongate. 31. Larva green. — Most nepticulid larvae are yellow or more transparent white. Bright green larvae occur scattered throughout the family, especially in Stigmella, but in Ectoedemia, apart from all species in the suberis group only algeriensis and gilvipen- nella have green larvae. Within the suberis group, aegilopidella takes an isolated position, but the remaining species VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia - 4 n u 535 o Sao mn 2 jeeo--soooooooooEo EM aa © ER = So: oc S | oo oo--000000000000 OU s x li Sn. mh®2|2o0o00--000-000000-.& > n ES pi J|E20200-0000-000000-.& Ss 3 vo 5.88 nb2)2o00--0000-000000-—-& 3 vo “Zé arc |EEEo--0000-000000- T vo wlooo--0000-0000o00-XK E à | atr + La) u DO ang ?[200—--0000-000000-m LD] BES hex S|000+-0000-0000000# (os) 39 -|oooo0-000000000oo-X% Som agr + SUS piQJ)e2o2o0-00000-000-00€& vu iene ery Q|-TOoo0o-0000000-000-.& EL i È Se BO os MAO SOS SS) SoS cS << [sci E > pe sp N [ma 0 a. 0.0 0.000: = = em A À À O n. oa 3 ce) ALES phy 2 |—-000-00c6c00----00 .Q PA SPINO SA | lie RETTO TOTO SIS) ©) GS) SS) (OP tal SL mn Inge Ss mo00-00000------o0% el ols ub2|-200-00000--=----00 2883 con Q Ooon-onr- O0 On — MA OC (OSS DO -|-o0o--0o00-0------o0 ZE | bs © Sa à cer © oo--o00-0-. = — — © C ANS Zod ab Sjors sel nig © mo0o--000-0-----000% Sn alin & oon-n-o0n-.00.--on.n.oo(0 n So 5 Inde SRI ZOO ©} SV io ua i GSO ii © © @ O/ „ SBS hr % |-eoe2o-0000-- - 00-0000 = gee leu 2 mn Onn On Om ORO ROA. On. 2 398 nlooo-0-00----0000 Ss Os gl À © 0 6 & quia oo--o000---=-o00000% Dan a Do @ fy ae aloooo---0000000000 BI aeg — ee and2|eooo0o---0-000000000% > OT = LS ad 2 sub 5 |2Poo0o--=-=-000000000% 2 ag sp No) De De De De nt De De De Ae One. À À © © À. Oa. ek © pia Bes car DJeEEEE-- Hm mo00000000% © SE E85 + -oo-o2000000--.00095 o 5 pre — _ ENS Ea Co Zoeloes | Oooo | | ew m! ~ . e klia|---00000000000000w Der =! Bs tur -[l[---00000000000000on = _— dd 43 © fol hno|l---00000000000000n ZI — . . gum | nt a|=---00000000-00000on ae 8 Qe ss ’ a Eig ue © CL ad ae Ta fe} Il = s 3 O JT OS SAS Qos BS BSS SAS Sr 52) O TARA À nm on À à À À À + + + 89 90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 form a tight group with the following apomor- phies: 32. T7 and 8 with many long tactile setae. — This apomorphy occurs in various distant- ly related species, feeding on evergreen Quercus. It could be an apomorphy for this part of the suberis group, and secondarily lost in andalusiae. 33. Ductus spermathecae with more than 312 convolutions. — Although the increase of the number of convolutions occurred sev- eral times in Nepticulidae, it is supposed that it is an apomorphy for this part of the suberis group. The remaining species of Ectoedemia s.str. most likely form a monophyletic unit, at least based on character 34. In fig. 2 character 28 is also an apomorphy but as a parallelism with the suberis group. The phylogeny within this part of the genus is still far from resolved, the charac- ters showing a very complicated pattern, but a tentative phylogeny is given in fig. 3. 34. Gnathos with central element divided into distal spatulate and basal serrate part. — The single, smooth central element is the generalised condition in Nepticulidae (Sco- ble, 1983). The “divided” gnathos only oc- curs in the subbimaculella group, exclud- ing the two species complexes and nıgros- parsella, and in the angulifasciella group, excluding the first three species. Since the structure is so uniform, it is most unlikely that it originated twice independently, therefore its absence in part of these groups must be explained by reversal or its presence by underlying synapomorphy. Alternatively this character is only an apo- morphy for species 20—26 and 42—48 to- gether, in which case the hypotheses of monophyly of the subbimaculella group and angulifasciella group both must be re- futed, but on parsimonious grounds I pre- fer the present solution. E. terebinthivora and the subbimaculella group are here regarded as sister-groups on the basis of the following: 35. Signa distinctly dissimilar in shape. — Here the dorsal signum is much longer than the ventral, it reaches almost into the vestibulum, and the shape of the posterior part is different from the other signum. A slightly similar situation occurs in spiraeae. The subbimaculella group is considered a mo- nophyletic entity on the basis of the next two characters: 36. Corpus bursae without pectinations. — Pectinations on the bursa belong to the ground-plan of Nepticulidae, their loss is therefore an apomorphy. The only other Ectoedemia species with this character, probably as a parallelism, is ıntimella. 37. T7 with a distinct row of setae along ante- rior margin of T8. — This row occurs in most species of the subbimaculella group but also in preisseckeri and erythrogenella. Probably it has secondarily evolved into a group of long setae similar to character 32 in the species algeriensis and leucothorax. A large part of this group shares the follow- ing character: - 38. Costal bristles present in male. — The in- terpretation of this character is open to doubt. In itself the presence of costal bris- tles belongs to the groundplan of Nepticu- . lidae. Costal bristles and the male hair- pencil are homologous structures; they al- ways occur more or less in the same posi- tion, and hair-pencil and costal bristles are mutually exclusive. The presence of a hair- pencil is regarded as part of the groundplan of Ectoedemia s.str., and in this case the presence of costal bristles can best be ex- plained as a reversal and therefore an apo- morphy within the subgenus. The alterna- tive explanation that these species retained the plesiomorphic condition implies the parallel development of a hair-pencil in many cases, in which case it could be based on an underlying apomorphy. The evi- dence here is not sufficient to eliminate this explanation entirely, but the presence of this character in a group of species, which also shares other attributes favours the re- versal interpretation at present. The remaining three species, quinquella, alge- riensis and gilvipennella, possess a hair-pencil, in this interpretation the plesiomorphic condi- tion of 38. They are tentatively placed as the sis- ter-group of the other species of the subbimacu- lella group with the following possible apomor- phy: Î | 39. Forewing with a pale discal spot in second half. — A distinct feature of quinquella and algeriensis. E. gilvipennella has a com- pletely pale forewing with some scattered dark scales, which can be explained as an enlargement of the white spots and hence as a further step in the transformation se- ries 39, but this remains a weak character which needs corroboration. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 91 E. quinquella and algeriensis clearly form a pair of sister species, based on the following apomorphy and corroborated by their high sim- ilarity: 40. Male hindwing with patch of special scales near hair-pencil. E. leucothorax cannot be placed with certain- ty in the cladogram. The next three species, ha- raldi, ilicis and heringella are considered to form a monophyletic group, merely based on simila- rities. Especially mine form and life history are very similar. E. ilicis and E. heringella are most likely sister-species with the following apomor- phy: 41. Loss of costal spot in forewing. E. nigrosparsella, the albifasciella complex and the subbimaculella complex form a well de- fined monophyletic entity based on the follow- ing characters: 42. Valva with many setae on inner (mesal) surface. 43. Gnathos smooth and undivided. — A re- versal of character 34. 44. Mine type. — The albifasciella and subbi- maculella complexes have a unique mine type: a narrow gallery, usually following a vein, abruptly enlarging ınto a square or triangular blotch, often in vein axil. Only nigrosparsella has a different type but is on other grounds regarded as a relative of the albifasciella complex. 45. Hostplant deciduous Quercus. — This might be correlated with character 44. Ev- ergreen species form the majority of the Fagaceae and deciduous forms occur ex- clusively in temperate regions. From this fact it seems likely that ancestral oak-min- ing Nepticulidae lived on evergreen oaks. However, this point needs further re- search, especially in the extensive ever- green cupuliferous forests in east and southeast Asia. The albifasciella complex and nigrosparsella are characterised by: 46. Convolutions of ductus spermathecae widened. — Except in albifasciella the number of convolutions is also increased. On the grounds of the number of convolu- tions nigrosparsella seems close to contor- ta, but otherwise it is quite different from the complex. The subbimaculella complex can be charac- terised by: 47. Forewing with basal spot. The angulifasciella group is a rather loose ag- gregrate of species sharing the hostplant (Rosa- ceae, character 48), which can hardly be re- garded as a sound synapomorphy, considering the wide variety of unrelated Nepticulidae and many other Microlepidoptera feeding on this plant family. The following morphological character might be the only true synapomorphy for this group: 49. Forewing with metallic coloured fascia. — This is not or hardly developed in spiraeae and hexapetalae, and remains a weak char- acter. It is not unlikely that this group is in fact paraphyletic in terms of either the sub- bimaculella or occultella group or both. Especially E. erythrogenella is different from other species in the group, and shares character 37 with the subbimaculella group, and moreover resembles albifasciel- la externally. E. spiraeae and agrimoniae form a pair on the following grounds: 50. Vaginal sclerite lost. 51. Spiculate pouch lost. 52. T8in female divided. The remaining species of this group possibly form a monophyletic entity based on their simi- larity, but I failed to find a distinct apomorphy. The branching within this group is presented as an unresolved trichotomy between hexapetalae, the angulifasciella complex and the pair spino- sella and mabalebella. For the angulifasciella complex the following character is an apomor- phy: 53. Carinae with many basal spines. The occultella group is a sound monophyletic entity on the following apomorphies: 54. Cilia-line lost. 55. Tegumen cuspidate. 56. Carinae divided into blunt ending process- es. 57. Pectinations on bursa arranged in bands. 58. Hostplant Betulaceae. However, the affinities of this group are not clear. It must be placed somewhere between the suberis and angulifasciella groups since it shares characters 27 and 28 with those groups, but any indication about its sister-group relationship is lacking, hence its tentative placement at the end. BIOGEOGRAPHY Discussion of the biogeography is limited to a few remarks owing to the scanty knowledge of the distribution. The subgenera discussed here are both widely distributed in the Holarctic re- gion, probably with the highest number of 92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 (mostly unknown) species in the Eastern Pal- aearctic, as indicated by some preliminary work on that fauna. In the Southern Hemisphere only Ectoedemia s. str. is known with three species from South Africa (Scoble, 1978, 1979), a very low number in relation to the total number of species (Scoble, 1983). In a large collection of Australian Nepticulidae the genus was not pre- sent (Scoble, 1983), neither was it in New Zea- land (Donner and Wilkinson, pers. comm.). Un- fortunately little is known from the Oriental and Neotropical regions, so that the conclusion that the group is predominantly Holarctic is not yet justified. In fact, the high number of Faga- ceae feeding species might lead to the assump- tion that these subgenera are well represented in the Fagaceous forests of the Oriental region. Many of the widespread European species probably have a distribution which goes much further east, but hardly any data are available from the Soviet Union. Many species are re- stricted to the mediterranean region and some of these (quinquella, erythrogenella) have an at- lantic-mediterranean distribution type. The species pairs heringella-ilicis and contor- ta-pubescivora are examples of vicariant species pairs with an eastern and western mediterranean element. These species are closely related and feed on the same hostplant, so that they most likely originated from populations isolated dur- ing the glaciation in west or east mediterranean refugia. ACKNOWLEDGEMENTS I would like to thank the following people for the loan of material and for information: Mrs. P. Arduino (Roma), Prof. Dr. R. Buvat (Mar- seille), Dr. A. Casale (Torino), Dr. D. R. Davis (Washington, D.C.), Mr. G. Derra (Bamberg), Dr. W. Dierl (Munchen), Dr. J. P. Duffels (Amsterdam), Dr. W. Foster (Cambridge), Mr. A. van Frankenhuyzen (Wageningen), Dr. M. Geisthardt (Wiesbaden), Dr. Ph. Georges (Brussel), Mr. C. Gielis (Lexmond), Dr. L. Gozmany (Budapest), Dr. F. Gregor (Brno), Mr. B. Gustafsson (Stockholm), Dr. H. Hanigk (Lobbach), Prof. Dr. H. J. Hannemann (Berlin), Mr. W. Hogenes (Amsterdam), Mr. K. J. Huis- man (Melissant), Mr. R. Johansson (Växjô), Dr. R. de Jong (Leiden), Mr. ©. Karsholt (Koben- havn), Dr. F. Kasy (Wien), Dr. J. Klimesch (Linz), Mr. J. Koster (Callantsoog), Dr. N. P. Kristensen (Kobenhavn), Mr. J. Kuchlein (Wa- geningen), Mr. J. Kyrki (Oulu), Mr. P. Leraut (Paris), Dr. G. Luquet (Paris), Dr. J. Matter (Strasbourg), Dr. E. S. Nielsen (Canberra), Prof. Dr. D. Povolny (Brno), Dr. R. Puplesis (Leningrad), Prof. Dr. J. Razowski (Krakow), Dr. U. Roesler (Karlsruhe), Dr. K. Sattler (London), Prof. Dr. W. Sauter (Zürich), Dr. H. Schröder (Frankfurt), Mr. W. Speidel (Karlsruhe), Dr. P. D. Syme (Salt Ste. Marie), Dr. E. Traugott-Olsen (Marbella), Mr. K. Tuck (London), Dr. P. Viette (Paris), Dr. S. E. White- bread (Magden) and Mr. J. Wolschrijn (Apel- doorn). Of these I would especially acknowl- edge the help of R. Johansson, for the wealth of unpublished data he allowed me to use in this work. For their hospitality and help during col- lecting trips I am indebted to Col. and Mrs. A. M. Emmet (Saffron-Walden), Mrs. A. Hallın and Mr. E. Traugott-Olsen (Marbella) and Dr. F. Kasy (Wien). I would like to thank my col- leagues of the Vrije Universiteit — Dr. Koos . Boomsma, Dr. Georgina Bryan and Dr. Steph Menken — for their advice and cooperation during the course of my studies and in joint col- lecting-trips, and Steph Menken also for all his data on allozyme studies of Ectoedemia. I am indebted for advice and critical remarks to Prof. Dr. C. Wilkinson, who also initiated this study. Similarly Dr. R. de Jong is acknowledged. Technical assistance by Mr. Kees Alders, Mr. Bart Jan van Cronenburg, Mrs. Daisy Kloos and Mr. Adri Rol, especially in preparation of genitalia and rearing work is much appreciated. I wish to acknowledge Messrs T. Feijen, P. W. A. van Huijstee and B. H. van Nifterik for as- sisting with photographic work, Mrs. Silvia Richter and Désirée Hoonhout for typing the manuscript, and Mr. L. Sanna for preparing figs. 1—3. For collecting trips to Greece and Central Europe grants were received from the Nether- lands Organisation for the Advancement of Pure Research (ZWO) and the Uyttenboogaart- Eliasen foundation. ‘REFERENCES Ballet Fletcher, W. H., 1882. Nepticula agrimoniae Heyden, a species new to Britain. — Entomolo- gist’s mon. Mag. 18: 211. Bedell, G., 1848. Description of Microsetia quinquel- la, a new species of moth of the family Tineidae. — Zoologist 6: 1986. Beirne, B. P., 1945. The male genitalia of the British Stigmellidae (Nepticulidae) (Lep.). — Proc. R. Ir. Acad. Be 50: 191—218. Borkowski, A., 1969. Studien an Stigmelliden (Lepi- doptera). Ti Zur Verbreitung, Biologie und Okologie der Stigmelliden in den polnischen Su- deten. — Polskie Pismo ent. 34: 95—122. 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Zur Kenntnis der Verbreitung der Nepticuliden in den Reichsgauen Wien und Niederdonau (Lepidopt.). — Z. wien. ent. Ges. 29: 3—6, 60—64, 78—91, 107—122. 98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Index to (sub)genera and species treated Reference to the first page of the treatment of each species is given only. Synonyms are given in italics, unavailable names provided with a double dagger (+) and misidentifications are cit- ed in square brackets. aegilopidella agrimomella agrimoniae agrimoniella albifasciella albifasciella-complex algeriensis algeriensis, cf + alliatae alnifoliae amani andalusiae angulifasciella angulıfasciella partim angulifasciella-complex angulifasciella-group apicella arcuata arcuatella arcuatella partim arcuosella argentipedella argentipedella partim argyropeza Herrich-Schäffer argyropeza Zeller [argyropeza sensu Beirne] [argyropeza sensu Petersen] [argyropeza sensu Stainton] argyropezella Doubleday argyropezella Herrich-Schäffer aterrima + aterrimoides atricolella atricollis atrifrontella bistrimaculella brunniella caradjai castaneae-group cerris contorta cursoriella Dechtiria Ectoedemia erythrogenella gilvella gilvipennella hannoverella haraldi heringella heringi heringiella hexapetalae hispanica + houzeaui + ilicella ilicis intimella + juncta klimeschi leucothorax liebwerdella liechtensteini liguricella lindquisti longicaudella mahalebella + malivora marionella mediofasciella [mediofasciella sensu Bradley] minimella minorella monemvasiae montissancti + morosella mucidella niculescui nigrociliella nigrosparsella nuristanica occultella occultella-group peinu phyllotomella populella populella-group Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia populi-albae preisseckeri preisseckeri-group prinophyllella + prunivora pubescivora quercifoliae quinquella rubivora [rubivora sensu Walsingham | sativella schleichiella simplicella “species” (specimen 1843) “species” (specimen 1375) “species” Gustafsson “species” Klimesch “species” van Nieukerken “species” Povolny & Gregor “species” Skala spinosella spiraeae spireae 31 57 37 staphyleae strigilella subapicella subbimaculella subbimaculella-complex subbimaculella-group suberis suberis-group terebinthivora terebinthivora-group Trifurcula partim turbidella Herrich-Schäffer turbidella Zeller turbulentella utensis viridella viridicola woolhopiella zimmermanni Zimmermannia 99 71 78 52 57 56 43 40 38 63 63 277 55 31 35 69 40 80 80 59 17 100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 N N Figs. 4—7. Schematic diagrams of genitalia in Ectoedemia (s.str.). 4,5, 3 genitalia, E. occultella; 4, ventral as- pect; 5, lateral aspect. a = aedeagus; ca = carinae; de = ductus ejaculatorius; g = gnathos; lv = lateral arm of vinculum; t = tegumen; tr = transtilla; v = valva: vp = ventral process of aedeagus; vv = ventral plate of vincu- lum. 6, 7, 2 genitalia. 6, E. albifasciella, ventral aspect; 7, E. hannoverella,lateral aspect. aa = anterior apo- physes; ap = anal papillae; bc = bursa copulatrix; ds = ductus spermathecae; pa = posterior apophyses; s = signum; s7 = segment 7, s8 = sternite 8; sp = spiculate pouch; t8/9 = tergite 8/9; v = vestibulum; vs = vaginal sclerite. Scales: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 101 Figs. 8—9. Venation in Ectoedemia. 8, E. (Zimmermannıa) atrıfrontella, 3, slide VU 252, Netherlands, Hilver- sum; 9, E. (s.str.) intimella, 3, veins labelled, f = fold (no vein), slide VU 196, Netherlands, Schinveld. Figs. 10—14. Hindwings of Ectoedemia (Zimmermannia), 3, showing hair-pencil and surrounding special scales. Normal hindwing scales and fringe not drawn. 10, E. atrifrontella, Netherlands, Hilversum; 11, E. liebwerdella, East Germany, Tharandt; 12, E. longicaudella, Netherlands, Nijmegen; 13, E. monemvasiae, holotype, hair- pencil spread out; 14, E. amani, Sweden, Stockholm. Scales: 0.5 mm. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, aFL. 1, 1985 102 un Og :€Z Sum | :07 um OI : IZ ‘61 ‘ZI Sum 7 :9] {wm oor #7 “CC ‘81 ‘GI ‘SOJES ‘pouad-1rey ‘D]jrsaaouuvg. “FT “47 ‘rep ‘wopr ‘eq ‘Jouad-aey 9779170290 ‘4 ‘77 ‘sayeos [Puad-IeY Jo soos ‘v]papnvduo] (viuunrusgumiz) ‘q ‘17 ‘61 WOT] 2JE9S 2UO JO | Te1aq ‘OZ ‘Saqeos [eroads JO [reep ‘wap] ‘61 ‘(SMOIE) safes jeroods pue puad-ırey yıım ‘pjasowids ‘7 ‘gy ‘s100dse [eszop ‘vrıu2p2097 © JO sainionss SUIMPUIH ‘47—g8] ‘s31] ‘190d5se jenu ‘saejo esse], ‘/] ‘249 punodwos jo grew q ‘91 ‘19adse [e1U0I1] “paAowsı Apıed sojess peoH “SI ‘S1][02143v vrwopaorg Jo A8ojoydiow ynpy "/1—G] ‘SSL aS) S D TS u © + © RQ TS x 8 -S Ss S IS S S D S À N MO) S © = 8 D 8 — N Q S = D + n = VAN NIEUKERKEN ‘unl ¢ :pe fumi OOI : LE “um 06 :0€ ‘67 ‘uN OZ ce ‘87 fumi | :97 ‘um Oy :97 ‘sofeag ‘ous oouıs Jo [reza] “pe f199dse Jessop “rppoguoufiuv (viuUvULaUUAZ) 7 ‘gg ‘avios aieunsod jo [e12Q ‘TE ‘81/2 UO avias suo] Burmoys ‘19adse [es10p ‘ovisvawauou (viuuvuinuuaz) “J “TE Sg audio = gi ‘/ audio = /1 ‘(sourds yim) g ortuaors = gs ‘/ u1aıs = /s ‘(6 903191 uo) aejjıded jeue = de aoadse Jessie] ‘wopr ‘oc ‘a0adse jezorejoszop “7773174220 °7 ‘67 ‘Uowopgeisod sfewa,g “p¢—67 ‘SBI,J ‘(mozre) seuLıes zo sourds [eseq Surmous ‘snsvapav pue avajea jo 11ed YIM Ioadse genua “poor “q ‘87 Sadadse [esoiey ‘vyjapuvaisuo] (vıuurwsowunz) ‘7 ‘47 eıfertuad ATEN ‘BELT 'SSIJ ‘2IMIONIIS a|eos BUIMOIOY “y7]997NII0 vIUIPIODT ‘9T—S7 ‘SÙ TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 104 ‘SOJEUI ur sojeos posijeioads sresıpur SMOIIVY -uop,WIEJNO ‘099010 ‘© Vyporrandig “FT ‘ch ‘adArojoy “9 ‘PAUVISISANU ‘FT ‘Tp “SION “uapamg “è wvv ‘7 ‘Ir "eiseAwouop ‘999915 ‘odAqered “è “arıspamauou ‘7 ‘op !adArojoy ‘9 ‘vo1uvdsiq -q Ge ÁqsBop ‘uapams ‘pP ‘vpapnonduo] “J gE !wapı ‘4 ‘vyjapsamqay "7 LE apuereyy, ‘Aueunon) ıseg ‘9 ‘vyjapsamgay '7 ‘9e ‘Buipey ospuerpon ‘spuepromon ‘9 ‘vyaruosfua ‘q ‘ge ‘dds (viuuvumsamunz) prwapaond "Ch —SE SLI 105 VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia -edung ‘à vlpram ‘J ‘0g :»dA10199] ‘9 ‘uagoasstaud 7 ‘Gy Teq ua Flog EIN © N E € fannT aq “SPUEJIOUION ‘© ‘vyjasaaouuvg J * ‘adArojoy “è ‘avisnjypuy ‘7 ‘75 ‘sauurd ‘ooueix ‘à ‘suaqns ‘7 TG ‘isoraoyA3en ‘AI ‘spueploygon ‘à ‘vzadout8ur ‘F ‘gp ‘ual “eLnsny ‘Pp “1qosawya 7 Lt ‘U9pIo] “spurts Spuejioyan ‘à ‘ajjaunui ‘7 ‘pp “dds (‚mss Gp -UINsIvUNIOG € ) DIUIPIOPA “TG hp ‘Sst TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 106 “elorensy ‘ureds “9 “sa ‘7 ‘19 ‘staegeuog ‘N ‘ureds ‘à pjvaog ‘4 ‘09 ‘ejjpgiem “ureds ‘adAavied à ‘x2409109n9] ‘7 ‘65 fauyeqyoior, “Aredunyg (mome) jouad-ırey xyoe]q Surmoys [re1op ‘9 vmauuadıana ‘7 ‘gg ‘931qaSeyna] we JOH ‘eimsny ‘© rpauuadıans ‘4 ‘15 :»dArojoy ‘à ‘sisuatsadyy ‘7 QG !fp/Asdurnopg “ureatig 1919) ‘à ‘vyanbumb ‘7 ‘sc !wapı ‘adAered “8 ‘vyapidojisav ‘4 ‘46 ‘sopoyy ‘999915 ‘adAiered “pamone sapeas ersods ‘p ‘yyyapidonsar ‘7 ‘eg ‘dds (‘ns's) vuuap20n7 "19— ES SS 107 tern Palaearctic Zimmermannia and Ectoedemia 5 We VAN NIEUKERKEN ‘puo aosagnd ae: -ofiujy ‘7 € 69:5 +9 en eo bs, si ~~ Sdf 7 Me Iezg ‘Are -9PIS1O pun € © ‘ç9 ‘apisioddn ‘9 OL ‘Sa]eos pasijeroads 9IEIIPUI C9 pur (9 UI SMOIIT ‘ad 100 Sun ‘à ‘524499 ‘7 ‘89 Suasutuase A ‘SPUEJIOUION ‘è 77] ‘79 Seyalry ‘erae[so8n x ‘2//23u1 aq “J +9 —79 ‘dnois 27]ajno U & ‘?7407U09 ‘TJ ‘OL onspfigjp "7 ‘29 seudopes ‘AMI è VUGGN c be Af ‘AP asuvdsoudiu ‘7 ‘9 dds (:ns:s ) VIUIP “eugapıeg ‘Aeıy “è 9 ‘adAiojoy “3 ‘arı 2094 “OLLI sd] TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 108 ‚sıoepng ‘Aresunp ‘à ‘avypradvxag ‘7 ‘6/ EIOAAG ‘299919 ‘9 ‘QDIUOWIABY “FJ ‘g/ SURIN “EIYPAOTSOYIEZD ‘ à ‘avavuds ‘q ‘// ‘puepiog ‘puejsug ‘© ‘yyauasorgidsa “7 ‘9/ :10oyq ‘299019 ‘4 ‘PAOUGIMQII “J “G/ “zemyg “UEI “è (GLET uauımads) dads ‘7 ‘p/ ‘adA10192] ‘© ‘npauo10]p(qd “7 “EL ‘B10gspyoeH "erusny ‘è Bumag “J ‘TL ‘oMnpPA 230H ‘spuryrayiany “è “vyaynovuaqans ‘q ‘14 ‘dds (mss) vrwopaoig ‘64-14 ‘SSI 109 Western Palaearctic Zimmermannia and Ectoedemia Van NIEUKERKEN ‘AEMION ‘9 “DIJIUIUIU “FJ ‘88 SuasuTUasE M ‘spueloyJaN ‘à ‘9772179290 ‘F -194]II0 ‘7 ‘98 ‘OUT, ‘Aye ay 9 rmogajprgru ‘7 ‘G8 ‘uadiny i “9 “vI]AIWNIAV “FJ ‘TS :OAO Xa SuIpaeiq ‘SpUe[JoUIN ‘è ‘s177091410 ‘7 € 18 :wns1eunog ‘SPUEJIIUION € 9 ‘ryjeusrfynèur 7 € 08 dds (ns “UdTePNAGIL ‘28 “Hopuory say -Aueu1n) ‘pamone sojeos jeroods ay Jo yoied “apısıapun ‘9 ‘pj SpUejJoUIoN ‘à ‘pyjasourds 7 ‘+8 :ylımasıaıur yy ‘spuellaygaN “Pp PAOMQNA 7 Cg !ersıdueı,] “999915, S) vU2p201 7 88—08 ‘Sr 110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 89—97. Ectoedemia spp., 6 genitalia (aedeagus removed in 93—95), ventral aspect. 89, E. atrifrontella, slide VU 087, Netherlands, Overveen; 90, E. liebwerdella, East Germany, Tharandt, slide on pin; 91, E. longı- caudella, slide VU 1835, Anatolia, Kizilcahamam; 92, E. hispanica, slide VU 1931, holotype; 93, E. monemva- siae, slide VU 1834, paratype, Anatolia, Kizilcahamam; 94, E. amani, slide MV 5752, Austria, Hundsheimer Berg; 95, E. nuristanica, slide MV 5402, holotype; 96, E. liguricella, slide VU 1828, Spain, Sierra Alfacar; 97. E. intimella, slide VU 1213, Netherlands, Rockanje. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 111 98 99 100 103 106 Figs. 98—106. Ectoedemia (s.str.) spp., & genitalia, ventral aspect (aedeagus removed in 103). 98, E. populella, slide VU 1252, syntype, USA; 99, E. hannoverella, slide MV 12202, West Germany, Baiern; 100, E. turbidella, slide MV 12206, Austria, Linz; 101, E. klimeschi, slide VU 1230, Austria, Linz; 102, E. preisseckeri, slide MV 12218, Austria, Wien; 103, E. caradjai, slide VU 1382, Hungary, Csopak; 104, E. spec. (specimen 1843), slide VU 1843, Spain, Rubielos de Mora; 105, E. suberis, slide VU 1112, France, “Nesp.”; 106, E. andalusiae, slide VU 1415, paratype, Spain, Camino de Ojen. 112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 107 108 ~ 109 110 1 114 115 Figs. 107—115. Ectoedemia (s.str.) spp., d genitalia, ventral aspect, aedeagus removed (except in 113). 107, E. aegilopidella, slide Klim. 1299, paratype, Greece, Rhodos; 108, E. quinquella, slide VU 869, England, Rainham; 109, E. cf. algeriensis, slide VU 1864, Morocco, Azrou; 110, E. gilvipennella, slide VU 1381, Hungary, Törökbálint; 111, E. leucothorax, slide VU 1885, paratype, Spain, Camino de Ojen; 112, E. haraldi, slide VU 868, paralectotype, France, Angouléme; 113, E. ilicis, slide VU 1420, Spain, Marbella; 114, E. heringella, slide VU 1395, Italy, Monti Aurunci; 115, E. heringella, slide RM 6666, Cyprus, Arakapos. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 113 116 117 118 119 121 122 123 124 Figs. 116—124. Ectoedemia (s.str.) spp., & genitalia, ventral aspect, aedeagus removed (except in 118, 120, 122). 116, E. nigrosparsella, slide VU 1378, Hungary, Törökbálint; 117, E. albifasciella, slide VU 864, Nether- lands, Hilversum; 118, E. cerris, slide VU 1729, Hungary, Szar; 119, E. pubescivora, slide VU 1342, paralecto- type, Switzerland, Somazzo; 120, E. cf. contorta, slide VU 909, Austria, Hundsheimer Berg; 121, E. subbimacu- lella, slide VU 863, Netherlands, Hilversum; 122, E. heringi, slide VU 1109, Poland, Bydgoszcz; 123, E. her- ingi, slide VU 867, paralectotype N. zimmermanni, Czechoslovakia, Libochowan; 124, E. liechtensteini, slide VU 1875, Hungary, Törökbalınt. 114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 EZ = 125 126 127 128 129 131 132 133 Figs. 125—133. Ectoedemia (s.str.) spp., & genitalia, ventral aspect, aedeagus removed in 127—129. 125, E. phyllotomella, slide VU 1849, Italy, Mte Vulture; 126, E. terebinthivora, slide VU 1250, Greece, Dhelfoi; 127, E. spiraeae, slide VU 873, Hungary, Sástó; 128, 129, E. erythrogenella, slide VU 946, lectotype, 129 focussed on more dorsal part; 130, E. hexapetalae, slide VU 1739, Hungary, Budaörs; 131, E. agrimoniae, slide VU 642, Greece, Evvoia; 132, E. agrimoniae, slide MV 12186, East Germany, Potsdam (focussed more dorsally); 133, E. angulifasciella, slide MV 12180, Austria, Hundsheimer Berg. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 115 136 134 135 139 137 138 140 141 ni 142 Figs. 134—142. Ectoedemia (s.str.) spp., d genitalia, ventral aspect. 134, E. atricollis, slide VU 1152, France, Clamart (from Crataegus); 135, E. rubivora, slide VU 1103, Denmark, Faaborg; 136, E. arcuatella, slide MV 12184, East Germany, Friedland; 137, 138, E. spinosella, slide VU 644, Greece, Arakhova (138 focussed more dorsally); 139, E. occultella, slide VU 1227, France, Pralognan; 140, 141, E. mahalebella, slide VU 997, Greece, Mt. Timfristos, (141 focussed more dorsally); 142, E. minimella, slide VU 1173, Norway, Rennebu. 116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 143—154. Ectoedemia (Zimmermannia) spp., ® genitalia. 143, E. atrifrontella, abdominal tip, slide VU 496, Netherlands, Nijmegen; 144, idem, bursa, slide VU 483, Netherlands, Hilversum; 145, 146, E. liebwerdel- la, slide VU 1873, East Germany, Tharandt; 147, 148, E. longicaudella, slide VU 860, Belgium, Aye; 149, 150, E. monemvasiae, slide VU 486, paratype, Greece, Monemvasia; 151, 152, E. amanı, slide VU 918, Sweden, Kul- laberg; 153, 154, E. nuristanica, slide MV 12141, paratype. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 117 156 158 160 162 164 166 Figs. 155—166. Ectoedemia spp., 2 genitalia. 155, 156, E. liguricella, slide VU 1414, Spain, Refugio de Juanar; 157, 158, E. intimella, slide VU 1254, England, Earls Colne; 159, 160, E. hannoverella, slide MV 12205, East Germany, Bautzen; 161, 162, E. turbidella, slide VU 1491, Netherlands, Santpoort; 163, 164, E. klimeschi, slide MV 12193, Austria, Hundsheimer Berg; 165, 166, E. argyropeza, slide VU 1933, Austria, Gumpoldskirchen. 118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 168 + Im. 172 174 - Figs. 167—175. Ectoedemia (s.str.) spp., 9 genitalia. 167, 168, E. preisseckeri, slide VU 1955, Hungary, Buda- | pest; 169, 170, E. suberis, slide VU 899, France, Golfe Juan; 171, 172, E. andalusiae, slide VU 1417, paratype, | Spain, Camino de Ojen; 173, E. caradjai, slide VU 1447, USSR, Babince, ex Quercus pubescens; 174, E. cf caradjai, slide VU 1867, Greece, Rhodos, ex Quercus infectoria; 175, E. cf caradjai, slide VU 1393, Greece, Rhodos, ex Quercus coccifera. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 119 178 r 179 180 181 182 183 184 Figs. 176—184. Ectoedemia (s.str.) spp., ® genitalia. 176, E. aegilopidella, slide VU 1390, paratype, Greece, Rhodos; 177, E. quinquella, slide VU 898, England, Rainham; 178, E. algeriensis, slide VU 1125, holotype; 179, E. gilvipennella, slide VU 1380, Hungary, Törökbalınt; 180, 181, E. leucothorax, slide Klim. 774, paratype, Spain, Marbella; 182, E. haraldi, slide VU 901, paralectotype, France, Angouléme; 183, E. ilicis, slide VU 943, paralectotype, Portugal, San Fiel; 184, E. cf. turbidella, slide VU 1492, Iran, Kered). 120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 186 188 nn 190 Figs. 185—196. Ectoedemia (s.str.) spp., © genitalia. 185, 186, E. heringella, slide VU 1398, Italy, Monti Aurunci; 186, 187, E. alnifoliae, slide RM 6572, holotype; 189, 190, E. nigrosparsella, slide VU 897, Italy, Sardegna; 191, 192, E. albifasciella, slide VU 892, Netherlands, Hilversum; 193, 194, E. cerris, slide VU 1333, lectotype; 195, 196, E. pubescivora, slide VU 1403, Italy, Sardegna, Belvi. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 121 200 202 203 © 205 Figs. 197—205. Ectoedemia (s.str.) spp., 2 genitalia. 197, 198, E. contorta, slide VU 1388, holotype; 199, 200, E. subbimaculella, slide VU 891, Netherlands, Hilversum; 201, 202, E. terebinthivora, slide VU 1245, Greece, Dhelfoi; 203, E. heringi, slide VU 894, paralectotype N. zimmermanni, Czechoslovakia, Libochowan; 204, E. hechtensteini, slide VU 1876, Hungary, Törökbálint; 205, E. phyllotomella, slide VU 1392, paralectotype, Italy, Ferrania. 122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 206 207 zo Figs. 206— 214. Ectoedemia (s.str.) spp., © genitalia. 206, E. spec. (specimen 1375), slide VU 1375, Iran, W. Shiraz; 207, E. erythrogenella, slide VU 972, France, Vannes; 208, E. spiraeae, VU 1868, paratype, Czechoslo- vakia, Cingov; 209, E. agrimoniae, slide VU 970, Greece, Kätsika; 210, E. hexapetalae, slide VU 1741, Hunga- ry, Budaörs; 211, E. angulifasaella, slide MV 12178, no locality, ex Rosa; 212, E. angulifasciella, slide VU 1345, paralectotype N. utensis, Switzerland, Zürich, ex Sanguisorba; 213, E. atricollis, slide MV 12177, Austria, Linz; 214, E. atricollis, slide VU 1186, Hungary, Budapest, ex Staphylea. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 123 215 es 216 ” 211 218 219 220 221 222 223 Figs. 215—222. Ectoedemia (s.str.) spp., 2 genitalia. 215, E. arcuatella, slide MV 12183, Austria, Wien; 216, E. rubivora, slide VU 964, Netherlands, Winterswijk; 217, E. spinosella, slide VU 947, lectotype; 218, E. spinosella, slide VU 1171, Greece, Arakhova, ex Prunus dulcis; 219, E. mahalebella, slide VU 976, Greece, Mt. Timfristos; 220, E. mahalebella, slide VU 1751, Hungary, Budaörs; 221, E. occultella, slide VU 1182, Austria; 222, E. mini- mella, slide VU 1220, France, Pralognan; 223, E. (Zimmermannia) amanı, detail of vestibulum with two groups of spines (arrows), slide VU 918, Sweden, Kullaberg. 124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 224230. Ectoedemia (s.str.) spp., details of 2 genitalia. 224, 225, 227, 228, Vestibulum; 226, 229, 230, Part of bursa with signa. 224, E. preisseckeri, with dense concentration of pectinations (arrow), slide VU 1955; 225, E. terebinthivora, spiculate pouch with single, long pointed spines, slide VU 1245; 226, E. agrimoniae, “spiny” signa, slide VU 970; 227, E. hannoverella, spiculate pouch with “single” spines, slide VU 1208; 228, E. contorta, spiculate pouch with single and grouped spines, slide VU 1388, holotype; 229, E. hannoverella, bursa with pectinations, slide VU 1208; 230, E. contorta, bursa smooth, slide VU 1388, holotype. 125 VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia ‘uredg ‘odAered ‘ogg, NA opijs ‘Yyjap4samgay A TET “Muy, ‘vnuvdsig ‘4 vez suayosryspjoduiny ‘ernsny “gEIZI AW apiJs ‘pyjopnva18uoy ‘1 “€¢T puereyr "wur 600 [EIS “EIO ap SOpeIqny fut BEG NA >PIIS ‘vyjaquosfisaw ‘7 ‘JET ‘20adse (zauur) jesıop ‘enger ‘9 ‘dds (pruuvrwusswunz) “Aueunan) 1seq ‘/6h] NA PIS VIMIPIONI “HETE SSI TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 126 “WU S0:0.:2je9g “woes essoIg ‘ureds ‘8781 NA 2PIS ‘27/224N87 “7 ‘SET ‘ad Aojoy ‘Tops AIN apts waruvastanu “gq ‘LET ‘Sinqnouipiso[y “eLnsny ‘6981 NA PIS Turm ‘7 ‘OEZ eiseaumuop ‘399219 ‘odArered 7/61 NA opijs ‘avisvawauow ‘7 ‘gez ‘29adse (tuur) [es1op ‘EATEA “9 “dds (mmuurwisswunz) viwapaord "SET—SET SSI gee VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 127 239 241 Figs. 239—244. Ectoedemia (s.str.) spp., populella and preisseckeri groups, 6 valva, dorsal (inner) aspect. 239, E. intimella, slide VU 1253, England, Earls Colne; 240, E. populella, slide VU 1252, syntype, USA; 241, E. han- noverella, slide MV 12202, West Germany, Baiern; 242, E. turbidella, slide MV 12206, Austria, Linz; 243, E. klimeschi, slide VU 1230, Austria, Linz; 244, E. preisseckeri, slide MV 12218, Austria, Wien. Scale: 0.05 mm. 128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 246 250 Figs. 245—250. Ectoedemia (s.str.) spp., suberis group, d, valva, dorsal (inner) aspect. 245, E. caradjai, slide MV 12153, Austria, Gumpoldskirchen; 246, E. cf. caradjai, slide Klim. 4200, Greece, Rhodos, from Quercus infectoria; 247, E. spec. (specimen 1843), slide VU 1843, Spain, Rubielos de Mora; 248, E. suberis, slide VU 1112, France, “Nesp.”; 249, E. andalusiae, slide VU 1416, paratype, Spain, Camino de Ojen; 250, E. aegilopi- della, slide Klim. 1299, paratype, Greece, Rhodos. Scale: 0.05 mm. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 129 254 Figs. 251—256. Ectoedemia (s.str.) spp., subbimaculella-group, 3, valva, dorsal (inner) aspect. 251, E. quin- quella, slide VU 1111, England, Rainham; 252, E. cf. algeriensis, slide VU 1864, Morocco, Azrou; 253, E. gilvi- pennella, slide VU 1381, Hungary, Törökbálint; 254, E. leucothorax, slide VU 1885, paratype, Spain, Camino de Ojen; 255, E. haraldi, slide VU 942, Portugal, [San Fiel]; 256, E. ilicis, slide VU 1358, lectotype. Scale: 0.05 mm. 130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 257262. Ectoedemia (s.str.) spp., subbimaculella group, &, valva, dorsal (inner) aspect. 257, E. ılıcıs, slide VU 1420, Spain, Marbella; 258, E. heringella, slide VU 1395, Italy, Monti Aurunci; 259, E. heringella, slide RM 6666, Cyprus, Arakapos; 260, E. nigrosparsella, slide VU 1378, Hungary, Törökbalınt; 261, E. albifasciella, slide VU 637, Netherlands, Winterswijk; 262, E. cerris, slide VU 1729, Hungary, Szar. Scale: 0.05 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 131 Figs. 263—268. Ectoedemia (s.str.) spp., subbimaculella and terebinthivora group, 6, valva, dorsal (inner) as- pect. 263, E. pubescivora, slide VU 1342, paralectotype, Switzerland, Somazzo; 264, E. cf. contorta, slide VU 1387, Hungary, Nagykovacsi; 265, E. subbimaculella, slide VU 1105, France, Alouette Pessac; 266, E. heringi, slide MV 12142, Austria, Klosterneuburg; 267, E. phyllotomella, slide Klim. 269, paralectotype, Italy, Ferrania; 268, E. terebinthivora, slide VU 1250, Greece, Dhelfoi. Scale: 0.05 mm. 132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 269 273 Figs. 269—274. Ectoedemia (s.str.) spp., angulifasciella group, 6, valva, dorsal (inner) aspect. 269, E. erythro- genella, slide VU 1170, paralectotype, France, Vannes; 270, E. agrimoniae, slide VU 642, Greece, Evvoia; 271, E. spiraeae, slide VU 873, Hungary, Sástó; 272, E. hexapetalae, slide VU 1739, Hungary, Budaörs; 273, E. angulifasciella, slide MV 12180, Austria, Hundsheimer Berg; 274, E. atricollis, slide VU 608, Netherlands, Winterswijk. Scale: 0.05 mm. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 133 276 soo C Cay C c € OMO, 6) o o (©) O £ < € 278 279 Figs. 275—280. Ectoedemia (s.str.) spp., angulifasciella and occultella group, &, valva, dorsal (inner) aspect. 275, E. arcuatella, slide MV 12184, East Germany, Friedland; 276, E. rubivora, slide VU 1103, Denmark, Faa- borg; 277, E. spinosella, slide VU 1137, Netherlands, Gulpen; 278, E. mahalebella, slide VU 881, Greece, Mt. Timfristos; 279, E. occultella, slide VU 1226, Netherlands, Rockanje; 280, E. minimella, slide VU 1173, Nor- way, Rennebu. Scale: 0.05 mm. 134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 | 281 283 D 284 285 286 287 288 289 290 291 292 Figs. 281—292. Ectoedemia spp., 6 genitalia, gnathos, ventral aspect (287 ventro-caudal aspect). 281, E. atrı- frontella, slide VU 937, France, Digne; 282, E. liebwerdella, slide VU 1832, France, St. Barnabé; 283, E. longı- caudella, slide VU 983, France, Digne; 284, E. hispanica, slide VU 1931, holotype; 285, E. monemvasiae, slide VU 1372, paratype, Greece, Monemvasia; 286, E. amani, slide MV 5752, Austria, Hundsheimer Berg; 287, E. nuristanica, slide MV 5402, holotype; 288, E. liguricella, slide VU 1828, Spain, Sierra Alfacar; 289, E. intimella, slide VU 1253, England, Earls Colne; 290, E. hannoverella, slide VU 292, West Germany, Regensburg; 291, E. turbidella, slide MV 12206, Austria, Linz; 292, E. klimeschi, slide VU 1230, Austria, Linz. Scale: 0.05 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 135 294 297 298 de 307 308 309 310 Figs. 293—310. Ectoedemia spp., 6 genitalia, gnathos, ventral aspect. 293, E. preisseckeri, slide MV 12218, Austria, Wien; 294, E. caradjai, slide RJ 946, Italy, Monti Aurunci (slightly squashed); 295, E. spec. (specimen 1843), slide VU 1843, Spain, Rubielos de Mora; 296, E. suberis, slide VU 1112, “Nesp.”; 297, E. andalusiae, slide VU 1416, paratype, Spain, Camino de Ojen; 298, E. aegilopidella, slide Klim. 1298, paratype, Greece, Rhodos; 299, E. quinquella, slide VU 1110, France, L’Etang-la-Ville; 300, E. cf algeriensis, slide VU 1864, Mo- rocco, Azrou; 301, E. gilvipennella, slide Klim. 272, lectotype; 302, E. leucothorax, slide VU 1885, paratype, Spain, Camino de Ojen; 303, E. haraldi, slide VU 868, paralectotype, France, Angoulème; 304, E. ilicis, slide VU 1420, Spain, Marbella; 305, E. heringella, slide VU 1395, Italy, Monti Aurunci; 306, E. heringella, slide RM 6666, Cyprus, Arakapos; 307, E. nigrosparsella, slide VU 1736, Hungary, Törökbálint; 308, E. albifasciella, slide VU 240, Netherlands, Hollandse Rading; 309, E. albifasciella, slide VU 1199, Netherlands, Overveen (po- sition of gnathos slightly different from 308); E. cerris, slide VU 1729, Hungary, Szar. Scale: 0.05 mm. 136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 311 | 312 315 316 317 318 319 TE CC IS 320 321 322 | 323 324 325 326 327 328 Figs. 311—328. Ectoedemia spp., 8 genitalia, gnathos, ventral aspect. 311, E. pubescivora, slide VU 1342, para- lectotype, Switzerland, Somazzo; 312, E. cf. contorta, slide VU 909, Austria, Hundsheimer Berg; 313, E. subbi- maculella, slide VU 1105, France, Alouette Pessac; 314, E. heringi, slide VU 1731, Hungary, Pécs Mecsek; 315, E. heringi, slide VU 867, paralectotype N. zimmermanni, Czechoslovakia, Libochowan; 316, E. terebinthivora slide VU 1250, Greece, Dhelfoi; 317, E. erythrogenella, slide VU 946, lectotype; 318, E. spiraeae, slide VU 1187, Hungary, Sástó; 319, E. agrimoniae, slide VU 642, Greece, Evvoia; 320, E. hexapetalae, slide VU 1740, Hungary, Budapest; 321, E. angulifasciella, slide VU 1870, Netherlands, Ootmarsum; 322, E. atricollis, slide VU 1152, France, Clamart; 323, E. arcuatella, slide MV 12184, East Germany, Friedland; 324, E. rubivora, slide VU 1001/1002, Netherlands, Winterswijk; 325, E. spinosella, slide VU 1139, paralectotype, France, Vannes; 326, E. mahalebella, slide VU 997, Greece, Mt. Timfristos; 327, E. occultella, slide VU 1495, Netherlands, Kerkrade; 328, E. minimella, slide VU 825, Norway, Rennebu. Scale: 0.05 mm. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 137 334 335 336 Figs. 329—336. Ectoedemia (Zimmermannia) spp-, d genitalia, capsule, ventral aspect, outline of left valva dot- ted or omitted. 329, E. atrifrontella, slide VU 937, France, Digne; 330, E. lebwerdella, slide Carolsfeld-Krause, East Germany, Tharandt; 331, E. longicaudella, slide VU 983, France, Digne; 332, E. hispanica, slide VU 1830, paratype, Spain, Rubielos de Mora; 333, E. monemvasiae, slide VU 476, paratype, Greece, Monemvasia; 334. E. amanı, slide VU 848, Sweden, Stockholm; 335, E. nuristanica, slide MV 5402, holotype; 336, E. liguricella, slide VU 929, France, “Nesp.”. Scale: 0.1 mm. 138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 337—345. Ectoedemia (Zimmermannia) spp., 6, aedeagus, ventral (337, 339, 341, 344) and dorsal aspect (338, 340, 342, 343, 345). 337, 338, E. atrifrontella, slide VU 087, Netherlands, Overveen; 339, 340, E. longicau- della, slide VU 830, Netherlands, Nijmegen; 341, 342, E. amani, slide MV 5752, Austria, Hundsheimer Berg; 343, E. liebwerdella, slide Carolsfeld-Krause, East Germany, Tharandt; 344, 345, E. hispanica, slide VU 1931, holotype. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 139 Figs. 346—350. Ectoedemia (Zimmermannia) spp., 6, aedeagus, left lateral aspect, in 346, 347 and 349 vesica extracted, in 348 and 350 omitted. 346, E. atrifrontella, slide MV 12134, Austria, Gumpoldskirchen; 347, E. longicaudella, slide VU 830, Netherlands, Nijmegen; 348, E. liebwerdella, slide Carolsfeld-Krause, East Ger- many, Tharandt; 349, E. monemvasiae, slide VU 482, paratype, Greece, Monemvasia; 350, E. amani, slide MV 5752, Austria, Hundsheimer Berg. Scale: 0.1 mm. 140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 354 Figs. 351—358. Ectoedemia (Zimmermannia) spp., d, aedeagus, 351, 353, 356, ventral aspect, 352, 354, 357, dorsal aspect; 355, 358, lateral aspect. 351, 352, E. monemvasiae, slide VU 470, paratype, Greece, Monemvasia; 353—355, E. nuristanica, slide MV 5402, holotype; 356—358, E. liguricella, slide MV 5415, Morocco, Ou- kaim’den. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 141 364 365 366 367 368 Figs. 359—368. Ectoedemia (s.str.) spp., 6, aedeagus, ventral aspect. 359, E. intimella, slide VU 1213, Nether- lands, Rockanje. 360, E. populella, slide VU 1252, syntype, USA; 361, E. hannoverella, slide VU 278, Nether- lands, Winterswijk; 362, E. turbidella, slide MV 12206, Austria, Linz; 363, E. klimeschi, slide VU 1230, Austria, Linz; 364, E. preisseckeri, slide MV 12214, lectotype; 365, E. caradjai, slide MV 12152, Austria, Hackelsberg; 366, E. spec. (specimen 1843), slide VU 1843, Spain, Rubielos de Mora; 367, E. suberis, slide VU 1112, France, “Nesp.”; 368, E. andalusiae, slide VU 1416, Spain, Camino de Ojen. Scale: 0.1 mm. 142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 369 370 371 372 373 374 375 376 377 378 Figs. 369—378. Ectoedemia (s.str.) spp., 6, aedeagus, ventral aspect. 369, E. aegilopidella, slide Klim. 4107, ho- lotype; 370, E. quinquella, slide VU 1111, England, Rainham; 371, E. cf. algeriensis, slide VU 1864, Morocco, Azrou; 372, E. gilvipennella, slide VU 1737, Hungary, Törökbalınt; 373, E. leucothorax, slide VU 1892, holo- type; 374, E. haraldi, slide VU 1116, France, Villenave d’Ornon, a, idem, tip of aedeagus of paralectotype, slide VU 868; 375, E. ilicis, slide VU 1358, lectotype; 376, E. heringella, slide BM 22604, France, Corsica; 377, E. heringella, slide RM 6666, Cyprus, Arakapos; 378, E. nigrosparsella, slide VU 1736, Hungary, Törökbálint. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 143 379 380 381 382 383 384 385 386 387 388 Figs. 379—388. Ectoedemia (s.str.) spp., 6, aedeagus, ventral aspect. 379, E. albifasciella, slide VU 1199, Neth- erlands, Overveen; 380. E. cerris, slide VU 1729, Hungary, Szar; 381, E. pubescivora, slide VU 1342, paralecto- type, Switzerland, Somazzo; 382, E. cf. contorta, slide VU 1387, Hungary, Nagykovacsi; 383, E. terebinthivora, slide VU 883, Greece, Dhelfoi; 384, E. subbimaculella, slide VU 863, Netherlands, Hilversum; 385, E. heringi, slide VU 1109, Poland, Bydgoszcz; 386, E. phyllotomella, slide Klim. 269, paralectotype, Italy, Ferrania; 387, E. erythrogenella, slide VU 1170, paralectotype, France, Vannes; 388, E. spiraeae, slide VU 1187, Hungary, Sástó. Scale: 0.1 mm. 144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 389 390 391 392 393 394 395 396 : 397 398 Figs. 389—398. Ectoedemia (s.str.) spp., 6, aedeagus, ventral aspect. 389, £. hexapetalae, slide VU 1739, Hun- gary, Budaors; 390, E. angulifasciella, slide VU 1157, France, Chaville; 391, E. atricollis, slide VU 1152, France, Clamart; 392, E. arcuatella, slide MV 12184, East Germany, Friedland; 393, E. rubivora, slide 1103, Denmark, Faaborg; 394, E. agrimoniae, slide VU 642, Greece, Evvoia; 395, E. spinosella, slide VU 1137, Netherlands, Gulpen; 396, E. mahalebella, slide VU 1750, Hungary, Budaörs; 397, E. occultella, slide VU 1226, Netherlands, Rockanje; 398, E. minimella, slide VU 1184, Norway, Grovudalen. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 145 400 399 403 Oe 405 406 Figs. 399—406. Ectoedemia (s.str.) spp., 6, aedeagus. 399, 401, right lateral aspect, 400, 402, 403, left lateral aspect; 399, E. hannoverella, Netherlands, Winterswijk; 400, 401, E. klimeschi, slide VU 801, Austria, Linz; 402, E. preisseckeri, slide MV 12214, lectotype; 403, E. hexapetalae, slide VU 1494, Hungary, Budaors; 404, E. hexapetalae, dorsal aspect, detail of spinose lobe, slide VU 1739, Hungary, Budaörs; 405, 406, Aedeagus with vesica, carinae omitted, ventral aspect; 405, E. occultella, slide VU 1226, Netherlands, Rockanje; 406, E. mini- mella, slide VU 1184, Norway, Grovudalen. Scales, 399—403: 0.1 mm, 404: 0.05 mm, 405—406: 0.05 mm. 146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 407 408 409 410 411 412 414 415 417 418 3 419 420 Figs. 407—420. Ectoedemia (s.str.) spp., details of genitalia. 407—414, d, Tegumen (pseuduncus), ventral as- pect. 407, E. caradjai, slide VU 1861, Anatolia, Kizilcahamam; 408, E. suberis, slide VU 1496, Italy, Sardegna, Mt. Istiddi; 409, E. andalusiae, slide VU 1416, paratype, Spain, Camino de Ojen; 410, E. aegilopidella, slide Kl. 1298, paralectotype, Greece, Rhodos; 411, E. quinquella, slide VU 1111, England, Rainham (slightly squashed); 412, E. terebinthivora, slide VU 1249, Greece, Dhelfoi; 413, E. rubivora, slide 1001, Netherlands, Winterswijk; 414, E. minimella, slide VU 825, Norway, Rennebu; 415, idem, lateral aspect; 416—418, ©, Ductus spermathe- cae. 416, E. albifasciella, slide VU 893, Greece, Palaiokastron; 417, E. subbimaculella, slide VU 638, England, Weeley; 418, E. heringi, slide VU 1900, Spain, San Pedro de Alcantara; 419, E. pubescivora, vaginal sclerite, ventral aspect, slide VU 1403, Italy, Sardegna, Belvi; 420, E. algeriensis, spiculate pouch, dorsal aspect, slide VU 1125, holotype. Scales: 407—415: 0.05 mm; 416—418: 0.1 mm; 419, 420: 0.05 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 147 424 Figs. 421—424. Female postabdomen of Ectoedemia (Zimmermannia) spp., dorsal aspect. Setal sockets only completely figured on one half of T8. 421, E. atrifrontella, slide VU 483, Netherlands, Hilversum; 422, E. lieb- werdella, slide VU 1873, East Germany, Tharandt; 423, E. longicaudella, slide VU 862, Belgium, Aye; 424, E. monemvasiae, slide VU 812, paratype, Greece, Monemvasia. Scale: 0.1 mm. 148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 425—428. Female postabdomen of Ectoedemia (Zimmermannia) spp., dorsal aspect. 425, E. amani, slide MV 1723 (slightly squashed), Yugoslavia, Skopje; 426, E. nuristanica, slide MV 12141, paratype, setal patch on T7 indicated by broken line, setal sockets only partly figured; 427, E. liguricella, slide BM 22669, Spain, Sierra Alfacar; 428, E. liguricella, aberrant specimen, slide MV 12140, Morocco, Oukaim’den. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 149 Figs. 429— 434, Female postabdomen of Ectoedemia (s.str.) spp., populella group. 429, E. intimella, slide VU 1254, England, Earls Colne; 430, E. hannoverella, slide MV 12205, East Germany, Bautzen; 431, E. turbidella, slide VU 1491, Netherlands, Santpoort; 432, E. cf. turbidella, slide VU 1492, Iran, Keredj; 433, E. klimeschi, slide VU 1231, Austria, Linz; 434, E. argyropeza, slide VU 1918, West Germany, Heidelberg. Scale: 0.1 mm. 150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 435—441. Female postabdomen of Ectoedemia (s.str.) spp., in 436 and 437 setal sockets only completed in right half of T8. 435, E. preisseckeri, slide MV 12215, paralectotype, Austria, Klosterneuburg; 436, E. caradjas, slide VU 1447, USSR, Babince; 437, E. suberis, slide VU 899, France, Golfe Juan; 438, E. andalusiae, slide VU 1417, paratype, Spain, Camino de Ojen; 439, E. aegilopidella, slide VU 1390, paratype, Greece, Rhodos; 440, E. quinquella, slide VU 898, England, Rainham; 441, E. algeriensis, slide VU 900, paratype, Algeria, Aures. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 151 Figs. 442—449. Female postabdomen of Ectoedemia (s.str.) spp., subbimaculella group. 442, E. spec. near alge- riensis, slide VU 1897, Spain, N. of Benahavis; 443, E. gilvipennella, slide VU 1380, Hungary, Torokbalint; 444, E. leucothorax, slide Klim. 774, paratype, Spain, Marbella; 445, E. haraldı, slide VU 901, paralectotype, France, Angouleme; 446, E. ilicis, slide VU 1352, France, “Nesp.”; 447, E. heringella, slide VU 902, paralectotype, Italy, Sicilia, Partinico; 448, E. heringella, slide RM 6667, Cyprus, Arakapos; 449, E. alnifoliae, slide RM 6572, holo- type. 448 and 449 slightly squashed. Scale: 0.1 mm. 152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 450—457. Female postabdomen of Ectoedemia (s.str.) spp., subbimaculella group. 450, E. nigrosparsella, slide VU 1379, Hungary, Törökbálint; 451, E. albifasciella, slide VU 892, Netherlands, Hilversum; 452, E. cer- ris, slide VU 1730, Hungary, Szar; 453, E. pubescivora, slide VU 1403, Italy, Sardegna, Belvi; 454, E. contorta, slide VU 1388, holotype; 455, E. subbimaculella, slide VU 891, Netherlands, Hilversum; 456, E. heringı, slide VU 895, paralectotype of N. zimmermanni, Czechoslovakia, Libochowan; 457, E. phyllotomella, slide VU 1392, paralectotype, Italy, Ferrania. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 153 Figs. 458—465. Female postabdomen of Ectoedemia (s.str.) spp. 458, E. spec. (specimen 1375), slide VU 1375, Iran, W. of Shiraz; 459, E. terebinthivora, slide VU 1248, Greece, Dhelfoi; 460, E. erythrogenella, slide BM 22675, France, Cannes; 461, E. spiraeae, slide VU 1868, paratype, Czechoslovakia, Cingov; 462, E. agrimoniae, slide VU 1136, Greece, Katsika; 463, E. hexapetalae, slide VU 1742, Hungary, Budapest; 464, E. angulifasciella, slide VU 969, Netherlands, Ootmarsum; 465, E. atricollis, slide VU 968, Netherlands, Ankeveense Plassen. Scale: 0.1 mm. 154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 466—471. Female postabdomen of Ectoedemia (s.str.) spp. 466, E. arcuatella, slide BM 22679, paralecto- type, Switzerland, Zürich; 467, E. rubivora, slide VU 649, Netherlands, Winterswijk; 468, E. spinosella, slide VU 947, lectotype; 469, E. mahalebella, slide VU 999, France, St. Thibaud-de-Couz; 470, E. occultella, slide VU 1183, Austria; 471, E. minimella, slide VU 1220, France, Pralognan. Scale: 0.1 mm. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 155 Figs. 472—479. Mines of Ectoedemia. 472, E. atrifrontella on Quercus robur (dried branch), Netherlands, Hol- landse Rading; 473, E. hebwerdella on Fagus sylvatica (dried bark), France, le Sappey-en-Chartreuse; 474, E. amanı on Ulmus sp., Sweden, from colour-slide R. Johansson; 475, E. hannoverella on Populus X canadensis, Netherlands, Bunde; 476, E. turbidella on Populus canescens, England, Loughton; 477, E. intimella on Salix cin- erea, Netherlands, Rockanje; 478, E. klimeschi on Populus alba, Austria, Linz; 479, E. preisseckeri on Ulmus sp., Austria, Wien. 156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 480—487. Mines of Ectoedemia on leaves of Quercus spp. 480, E. suberis on Q. suber, Spain, Sierra Blan- ca; 481, E. suberis or andalusiae on Q. coccifera, Spain, Marbella; 482, E. aegilopidella on Q. macrolepis, Greece, Rhodos; 483, E. caradjai on Q. pubescens, Greece, Oiti Oros; 484, E. algeriensis on Q. rotundifolia, Algeria, Arris; 485, E. quinquella on Q. robur, England, Herringswell; 486, E. gilvipennella on Q. cerris, Austria, Loret- to; 487, E. haraldi on Q. ilex, France, Roquefort. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 157 Figs. 488—495. Mines of Ectoedemia on leaves ot Quercus spp. 488, 489, E. ilicis on Q. suber, Spain, Sierra Blanca; 490, E. albifasciella on Q. petraea, West Germany, Wiesbaum; 491, E. nigrosparsella on Q. pubescens, Austria, Gumpoldskirchen; 492, E. cerris on Q. cerris, Austria, Hof am Leithagebirge; 493, E. pubescivora on Q. pubescens, Italy, Picinisco; 494, E. subbimaculella on Q. petraea, Yugoslavia, Bihac; 495, E. heringi on Q. petraea, Hungary, Törökbálint. 158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 496—504. Mines of Ectoedemia. 496, E. liechtensteini on Quercus cerris, Yugoslavia, Han Knezica; 497, E. phyllotomella on Q. cerris, Italy, Ferrania; 498, E. terebinthivora on Pistacia terebinthus, Greece, Dhelfoi; 499, E. erythrogenella on Rubus ulmifolius, Yugoslavia, Piran; 500, E. spiraeae on Spiraea media, Czechoslova- kia, Cingov; 501, E. agrimoniae on Agrımonia eupatoria, Austria, Hundsheimer Berg; 502, E. hexapetalae on Filipendula vulgaris, Austria, Gramatneusiedl; 503, E. angulifasciella on Rosa canina, Netherlands, Ootmarsum; 504, E. atricollis on Prunus avium, Austria, Hof am Leithagebirge. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 159 Figs. 505—513. Mines ot Ectoedemia. 505, E. atricollis on Staphylea pinnata, Austria, Hundsheimer Berg; 506, E. arcuatella on Fragaria vesca, Austria, Völkermarkt; 507, E. rubivora on Rubus saxatilis, Italy, Trento; 508, E. spmosella on Prunus spinosa, Netherlands, Gulpen; 509, E. spinosella on Prunus dulcis, Greece, Arakhova; 510, E. mahalebella on Prunus mahaleb, Yugoslavia, Selce; 511, E. occultella on Betula pubescens, Italy, Trento; 512, E. occultella on B. pendula, Austria, Nassfeld Pass; 513, E. minimella on B. pubescens, Norway, Rennebu. 160 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 514—519. Distribution maps of Ectoedemia spp. 514, E. atrifrontella; 515, E. liebwerdella; 516, E. longi- caudella; 517, E. hispanica (rectangles) and E. amani (dots); 518, E. hannoverella; 519, E. turbidella. VAN NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 161 Figs. 520—525. Distribution maps of Ectoedemia spp. 520, E. intimella; 521, E. argyropeza; 522, E. albifasciel- la; 523, E. subbimaculella; 524, E. heringi; 525, E. liechtensteini (dots) and E. phyllotomella (triangles). 162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 526—531. Distribution maps of Ectoedemia spp. 526, E. caradjai (dots) and E. andalusiae (triangles); 527, E. quinquella (dots), E. algeriensis and 6 cf. algeriensis (rectangles) and E. leucothorax and ® cf. algeriensis (tri- angle); 528, E. erythrogenella; 529, E. agrimoniae; 530, E. occultella; 531, E. minimella. Van NIEUKERKEN: Western Palaearctic Zimmermannia and Ectoedemia 163 (Figs. 532—537. Distribution maps of Ectoedemia ssp. 532, E. angulifasciella; 533, E. atricollis; 534, E. arcuatel- la; 535, E. rubivora; 536, E. spinosella; 537, E. mahalebella. 164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 1, 1985 Figs. 538—549. Distribution maps of Ectoedemia spp. 538, E. monemvasiae; 539, E. liguricella; 540, E. preis- seckeri (dots) and E. terebinthivora (rectangles); 541, E. klimeschi; 542, E. suberis; 543, E. gilvipennella (rectan- gles) and E. ilicis (dots); 544, E. heringella; 545, E. haraldi; 546, E. nigrosparsella (dots), E. aegilopidella (trian- gle) and E. alnifoliae (star); 547, E. pubescivora (dots) and E. contorta (rectangles); 548, E. cerris; 549, E. spi- raeae (dots) and E. hexapetalae (rectangles). sf RES A FAR i DEEL 128 AFLEVERING 2 1985 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING ser INHOUD M. R. DE Jonc. — Taxonomy and biogeography of Oriental Prasiini I: The genus Prasia Stal, 1863 (Homoptera, Tibicinidae), pp. 165—191. figs. 1—91. Tijdschrift voor Entomologie, deel 128, afl. 2 Gepubliceerd 20-X11-1985 î AE PRET. MARE tl ad TAXONOMY AND BIOGEOGRAPHY OF ORIENTAL PRASIINI 1: THE GENUS PRASIA STAL, 1863 (HOMOPTERA, TIBICINIDAE) by M. R. DE JONG Institute of Taxonomic Zoology (Zoölogisch Museum), University of Amsterdam, Amsterdam, The Netherlands ABSTRACT The taxonomic concept of the genus Prasia is re-established on account of characters found in the male genitalia. The genus now consists of a monophyletic group of seven spe- cies: P. faticina Stal, 1863, P. princeps Distant, 1888, P. breddini n. sp., P. sarasinorum n. sp., P. senilirata n. sp. and P. tuberculata n. sp., all from Sulawesi, and P. nigropercula n. sp. from the nearby Muna Island. All species are (re)described and structures of taxonomic im- portance as well as the whole insects are depicted. A key to the males is presented. Study of (type-)material established the new synonymy of P. culta Distant, 1898, with P. faticina. P. hariola Stal, 1863, and P. tincta Distant, 1909, are transferred to other genera of the Pra- siini. INTRODUCTION The present study of the genus Prasia Stal, 1863, is a further contribution to a revision of the tribe Prasiini, started by De Jong & Duffels (1981) and De Jong (1982). Preliminary phylo- genetic investigations plead in favour of com- mon ancestry of four Oriental genera belonging to this tribe: Prasia, Lembeja Distant, 1892, Ar- faka Distant, 1905, and Jacatra Distant, 1905. It is questionable whether the other genera placed in the Prasuni, viz. Lacetas Karsch, 1890, Irua- na Distant, 1905 (both from Africa), and Sapan- tanga Distant, 1905 (from South America), do form a monophyletic group together with the four above mentioned genera. Taxonomic and phylogenetic studies of the Oriental genera are to provide a basis for a reconstruction of their distributional history. The tribe Prasiini is presently distributed, as far as its Oriental members are concerned, in the Philippines, Sanghir, Sulawesi, Java, Lesser Sunda Islands, Misool, New Guinea and North- ern Queensland (Australia), and therefore con- sidered an excellent group to test Duffels’ idea about the role of Tertiary island arcs in the de- ‘velopment of the Cicadoidea fauna of Sulawesi, Moluccas, New Guinea and West Pacific. Duf- fels (1983a, b) indicated two routes of dispersal: in southern direction from the Philippines to Sulawesi, and in eastern direction from the Phil- ippines to the Moluccas, New Guinea and the 165 South-West Pacific. The eastern route itself is to be divided in two subroutes: the Inner- and Outer Melanesian Arcs (for a more detailed de- scription the reader is referred to Holloway (1979) and Duffels (1983a)). Study of the Prasii- ni will extend the area dealt with by Duffels to the Lesser Sunda Islands and Java, while materi- al obtained recently contains representatives from northern Borneo and Sumatera. The present study of the genus Prasia reveals that this very homogenous genus is confined to Sulawesi and the nearby Muna Island. HISTORY OF THE GENUS The genus Prasia Stal, 1863, was described to accommodate two new species, Prasia faticina Stal, 1863, and Prasia hariola Stal, 1863. How- ever, Stal (1862) had already referred Cephalo- xys foliata Walker, 1858, to Prasia, thus creating Prasia by indication (International Code of Zoological Nomenclature, 1985, Chapter iv, Article 12b, 5). For various reasons!) I favour suppression of this indication. A request to the International Commission is in preperation. !) According to the Rules (International Code of Zoological Nomenclature, 1985, Chapter iv, Article 12b 5), Cephaloxys foliata Walker, 1858, is the type- species by monotypy of Prasia Stal, 1862. Since P. fo- liata is currently placed in Lembeja Distant, 1892 (type-species L. maculosa (Distant, 1883)) by Distant (continued overleaf) 166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Pending the Commission’s decision I adhere to common usage (e.g. Distant, 1905) in using Pra- sia faticina as the type species of Prasia. ET Stal (1870) added P. fatiloqua Stal, 1870. In later publications P. princeps Dis- tant, 1888, P. culta Distant, 1898, P. tincta Dis- tant, 1909, and P. vitticollis Ashton, 1912, were described. The last mentioned species was transferred recently (De Jong, 1982) to the ge- nus Lembeja, on account of similarities in the opercula, male genitalia and wing-venation with some New Guinean representatives of this ge- nus. Furthermore some non-Prasiini have been (erroneously) attributed to the genus Prasia. Metcalf (1963) listed the following species under Prasia: P. culta, P. faticina, P. hariola, P. princeps and P. tincta. P. fatiloqua and P. fo- liata were listed as members of Lembeja, fol- lowing Horvath (1913) and Myers (1928, 1929) for fatiloqua and Distant (1906) for foliata. My study of the genera Prasia and Lembeja revealed that generic characters for Lembeja, as defined in the rudimentary vein (Distant, 1892) and the fusion of the Cu, and A, veins in the wing (Karsch, 1890b; Jacobi, 1903; Schmidt, 1925), were unsatisfactory. Breddin (1901) even ignored the existence of Lembeja and attributed all species studied by him and known at the time to Prasia. On the other hand he created the new genus Drepanopsaltria Breddin, 1901, for two Prasia species, viz. P. culta and P. princeps. Dis- tant (1905) separated Prasia and Lembeja on ac- count of characters in the tegmina. In Prasia the 3rd ulnar area is much shorter than the 1st, whilst in Lembeja they are more or less of equal length; the 4th ulnar area is much shorter than the radial area in Prasia, whilst in Lembeja the 4th is about as long or even just a little longer than the radial one. Apart from some borderline cases this is more or less correct, but one must bear in mind, that these characters are only valid when comparing Prasia with Lembeja. Here, a new concept for Prasia will be pre- sented, readily distinguishing Prasia from the other genera of the Oriental Prasiini. (1906) and other authors, all species of Lembeja should be transferred to Prasia, so that Lembeja be- comes a junior subjective synonym of Prasia. All authors have taken P. faticina as the type-spe- cies of Prasia. A valid name for Prasia in its common usage, 1.e. the concept based on P. faticina, could then be Drepanopsaltria Breddin, 1901 (type-species Prasia culta Distant, 1898, a junior synonym of Prasia fati- cina Stal, 1863). The name Drepanopsaltria has not been used as a valid name after 1905. A NEW CONCEPT FOR THE GENUS PRASIA Monophyly of the genus Prasia In my opinion P. faticina (with P. culta as a new synonym), P. princeps, and five new spe- cies, viz. P. breddini n. sp., P. nigropercula n. sp., P. sarasinorum n. sp., P. senilirata n. sp. and P. tuberculata n. sp., form a monophyletic group. At first sight the Prasia species can be sepa- rated from all other Oriental Prasiini by the straight margins of the pronotum collar (in lateral view), which are downgraded in the other genera. However, not much phylogenetic value can be attributed to this feature, since it is found in several other groups of Cicadoidea. The monophyly of Prasia in its new concept finds its justification in the supposed apomor- phy ou | in the degrading lateral margins of the pygofer between the lateral lobes and the’ caudal dorsal beak (fig. 8). Consequences of the new genus concept for remaining Prasia species Beside the species included now in Prasia, Metcalf (1963) listed two other species: P. hari- ola and P. tincta. As a consequence of the new concept for Prasia these species are transferred to other genera. P. hariola, of which recently a small, but very fine, series from Misoöl was discovered in the Vienna Museum, does not show the shape of the pygofer and the uplifted pronotum collar that characterize Prasia within the Oriental Prasiini. Moreover, P. hariola shows a structure of the male genitalia and a venation of tegmina and wings that characterize the previously monoty- pic genus Arfaka. This is ample evidence that P. hariola is to be considered a member of Arfa- ka (together with A. fulva (Walker, 1868)). A. hariola is described from the island of Misool and also known from New Guinea (Vogelkop) (Distant, 1892). P. tincta is transferred to the genus Lembeja, as it shows affinities with one of the four spe- cies-groups of Lembeja, the so-called L. fatilo- qua group. This group is characterized by the longitudinally medially dinted abdominal tergite 1 in the males (see also De Jong, 1982). L. tincta has been described from Bua-Kraeng and is now also known from Lompobattang (both lo- calities in South Sulawesi). MATERIAL AND METHODS The material examined for this study belongs DE Jone: Oriental Prasia 167 to 13 institutions, which are listed as deposito- ries. All type-material has been studied. Some museums have been visited in order to look for additional material. For tracing the localities I have used the Atlas van Tropisch Nederland (1938), the Times Atlas of the world (1973) and personal information from Dr. J. P. Duffels, Amsterdam. The distri- butions are presented on fig. 7 Terminology follows Duffels (1977, 1983a) with a few new morphological terms and modi- fications introduced, which will be explained in figs. 1—6, 8, 9, 11. The methods of Duffels (1977) are used for the examination of the male genitalia. Measurements were, apart from using a marking gauge with nonius, taken through a stereoscopic microscope with a specially de- signed ocular. DEPOSITORIES The abbreviations given below have been used in the lists of material and throughout the text. AMS Australian Museum, Sydney BIN Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussel BM British Museum (Natural History), London CNMW Naturhistorisches Museum, Wien DEI Deutsches Entomologisches Insti- tut, Eberswalde MHNG Muséum d’Histoire Naturelle, Gen- eve MNP Museum National d'Histoire Natu- relle, Paris MSNG Museo Civico di Storia Naturale “G. Doria”, Genova _MZB Museum Zoologicum Bogoriense, Bogor NRS Naturhistoriska Riksmuseet, Stock- holm 4 SMD Staatliches Museum für Tierkunde, Dresden TMB Termeszettudomany Muzeum, Bu- | dapest | ZMA Instituut voor Taxonomische Zo- dlogie, Zoölogisch Museum Amsterdam. ACKNOWLEDGEMENTS | Tam very obliged to Dr. J. P. Duffels and Dr. P. Oosterbroek for their critical reading of the manuscript. The help and hospitality of Dr. W. J. Knight, Mr. M. D. Webb and Mr. P. S. Broomtield (BM), Mr. R. Detry (BIN), Dr. A. Kaltenbach c.s. (CNMW), Dr. B. Hauser (MHNG), Dr. M. Boulard (MNP), Dr. R. Poggi (MSNG) and Dr. P. Lindskog (NRS), met with during my stay at their institutions is gratefully acknowledged. I feel also obliged to these persons for the loan of material under their care. The remaining materi- al was gratefully received from Dr. C. N. Smithers (AMS); Prof. Dr. H. J. Muller, Dr. G. Petersen and Dr. A. Täger (DEI); Dr. S. Ad- isoemarto (MZB); Dr. R. Emmrich (SMD); Dr. Z. Kaszab (TMB). I am indebted to Mr. G. Verlaan and Mr. J. Zaagman for technical assistance, to Mr. L. Van der Laan for the photographs, to Miss A. Stoel for typing the manuscript and to Dr. W. N. EI- lis for help with nomenclatural problems. The investigations werd supported by the Foundation for Fundamental Biological Re- search (BION), which is subsidized by the Netherlands Organization for the Advancement of Pure Research (ZWO). The participation of Dr. J. P. Duffels in the “Project Wallace” Expedition was supported by the Netherlands Foundation for the Advance- ment of Tropical Research (WOTRO). TAXONOMY Prasia Stal, 1863 Prasia Stal, 1862: 483 (num. nud.); Stal, 1863: 574 (original description); Walker, 1868: 94; Stal, 1870: 718; Distant, 1892: 103, 145; Breddin, 1901: 113, 153, 183, 200, 201; Jacobi, 1903: 12, 13; Dis- tant, 1905: 275, 278 (equals Drepanopsaltria Bredd.); Distant, 1906: 182, 183 (equals Drepa- nopsaltria Bredd.); Ashton, 1912: 221; Kato, 1932: 14, 30, 188, 189; Kato, 1956: 23, 70, 77, 78, 80; Metcalf, 1963: 423; Boulard, 1975: 315; Duf- fels, 1977: 205; Holloway, 1979: 235; De Jong & Duffels, 1981: 53, 61; De Jong, 1982: 182, 183; Duffels, 1983b: 492; Duffels & Van der Laan, 1985: 298. Drepanopsaltria; Breddin, 1901: 113, 183, 200, 201; Jacobi, 1903: 12; Metcalf, 1963: 425 (in synonymy of Prasia Stal). For a more complete list, refer to Metcalf (1963). Type-species: Prasia faticina Stal, designated by Distant, 1905: 278. Genital characters are given only for the males as there were too few females available to justify damaging specimens by dissection. When 168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 more material will be available the study of the female specimens will be continued. Diagnosis. Postclypeus obconically protruding in dorsal view. Antennal segment 1 extending clearly from under supra-antennal plate. Head 1.49— 2.07 X as long as width of vertex between eyes. Width of head 2.26—2.76 X width of vertex be- tween eyes. Head and pronotum together somewhat longer than meso- and metanotum together. Pronotum collar nearly twice as broad as width of head including eyes, not bent ven- trally, but forming a straight line along the lateral margins of the pronotum to the head (fig. 5) Fore femora with three thorns, basal cylindri- cal one with a dark brown apex; two apical thorns laterally compressed, middle thorn being almost as long as basal one, most apical one very small (fig. 3). Male opercula sickle-shaped and reaching 2nd sternite of abdomen. Female opercula small and more or less rounded. Figs. 1—5. Figures of Prasia: 1, male abdomen in ventral view, Prasia princeps; 2, right tegmen and wing, P. princeps; 3, male femur in lateral view, P. princeps; 4, aedeagus in lateral view (after treatment with 10% KOH), P. princeps; 5, head and pronotum in lateral view, P. princeps. (a 1—8 = 1st—8th apical area; A 1—3 = 1st—3rd anal vein; aap = aedeagal appendage process; as 1—2 = antennal segment 1—2; at = apical thorn; bt = basal thorn; Cu 2 = 2nd cubital vein; daa = dorsal aedeagal appendage; mop = male operculum; mpc = margin pro- notum collar; mt = middle thorn; rad a = radial area; sap = supra antennal plate; st 1,8 = Ist, 8th sternite; ua 1—4 = 1st—4th ulnar area.) DE Jone: Oriental Prasia pere — | r € PS pe ‘ Fig. 6. Tymbal, lateral view, P. nigropercula (Ir = long ridge, sr = short ridge, tr = tymbal ridge). Tegmina of males pale-hyaline, of females subhyaline. Wings pale-hyaline. Tegmina with fairly long apical areas, 3rd apical area less than 0.78 X the 4th one, 3rd ulnar area less than 0.87 X the Ist one, 4th ulnar area less than 0.77 x radial area. A, and Cu, veins fused up to the tegmen border. Anal field of wing enclosed by a fusion of the Cu, and A, veins or not. Anal lobe (area be- tween A, and A) apically fairly narrow (fig. 2). Male abdomen inflated, broadly raised me- dially along its whole length (tergite 2—8), but not carinate; ventrally bulged out. Tymbal ridges on tergite 2 distinct (fig. 6). Abdominal sternite 1 in males very large and swollen (fig. 1); its distal margin convex or slightly concave. Male sternite 8 apically slightly to strongly pointed. Tymbals with 6—9 (in P. princeps sometimes : 10) long ridges alternating with usually the same _ number of short ridges. Female abdomen distally more or less clubshaped, since greatest width of segment 9 is broader than width of hind margin of segment 8. Ovipositor sheath just or not reaching be- yond caudal dorsal beak. Lateral lobes of male pygofer fairly short, i swollen and acutely pointed, not extending be- | yond anal valves. Caudal dorsal beak laterally i compressed, and therefore very slender and long. Hind part of lateral margin between cau- dal dorsal beak and each of lateral lobes degrad- ing as in fig. 8. Claspers more or less swollen 169 and mostly elongate apically. Median uncus part above aedeagus small. Aedeagus slender, with two rounded, sometimes slightly dentate, lobes. Aedeagus with a dorsal appendage, splitting in two slender, apically pointed or rounded, pro- cesses, weakly sclerotized (fig. 4). Adjustment of aedeagus usually halfway the length of py- gofer. KEY TO THE MALES OF PRASIA 1. Operculum nearly black, contrasting with pale underside of the thorax. Muna........ nigropercula (p. 180) — Operculum concolorous with the (mostly pale) underside of the thorax 2. Medium-sized to large species (body length: 24.6—29.7 mm). Body colour green to greenish-olivaceous. Cu, and A, veins in the wing may be fused. North and Central Sulawesi sn Pain a Re er ded ar 3 — Small to medium-sized species (body length: 19.6—23.8 mm). Body colour brown, pale-brown or orange-brown. Cu, and A, veins never fused. Central, East and South Sulawesi a). a ne Den 4 3. Genitalia as in figs. 60—68. Cu, and A, veins usually fused. 9 (sometimes 8 or 10) long ridges alternating with usually the same number of short ridges. North Sula- WIESE le STE princeps (p. 182) — Genitalia as in figs. 76—81. Cu, and A, not fused. 6 long ridges alternating with the same number of short ridges. Central Sula- WIESE N | senilirata (p. 186) 4. Central fascia on the pronotum usually concolorous with it. Central and East Sula- WESTEN Mirth a Neen or la ER DR 5 — Central fascia usually dark-brown col- oured, or traces of such a coloration pre- sent. Central and South Sulawesi........ 6 5. Operculum rounded apically (fig. 39). Cen- traliSulawes Reese sarasinorum (p. 176) — ©Operculum pointed apically (fig. 14). East Sulawesi Ro e tuberculata (p. 178) 6. Genitalia as in figs. 8—13. South Sulawesi . . faticana (p. 170) — Genitalia as in figs. 21—26. Central Sula- WESTIE N fii en A ki breddini (p. 174) The females are not included in the key, since these are known with certainty of two species only. The females P. princeps are generally green to greenish-olivaceous (sometimes with a brownish tinge), whilst those of P. faticina are usually orange-brownish with olivaceous 170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 | tinges. Females of P. princeps (25.9—30.2 mm) Prasia faticina Stal, 1863 are usually larger than those of P. faticina (figs. 7—20, 86, 87) (24.6—26.2 mm). Third ulnar area of P. faticina Prasia faticina Stal, 1863: 574 Stal, 1870: 94; Distant, females often, but not always, shaped as in fig. 1888: 325; Distant, 1892: xiv, 145 (partim: only 20, of P. princeps females usually as in fig. 2, Makassar), pl. 7 figs. 14, 14a, b: 146; Jacobi, 1903: some females have a 3rd ulnar area of interme- 12; Distant, 1905: 278; Distant, 1906: 183; Kato, Alena shape (fig. 70). 1932: 189; Metcalf, 1963: 424. @ princeps O senilirata A faticina A breddini M sarasinorum O tuberculata y nigropercula 120° 1222 124° Fig. 7. Distributions of the species of Prasia, P. breddini, P. faticina, P. nigropercula, P. princeps, P. sarasino- rum, P. senilirata and P. tuberculata. DE Jong: Oriental Prasia 171 Prasia culta Distant, 1898: 97; Distant, 1905: 279 (equals Drepanopsaltria (Lembeja) culta Bredd.); Distant, 1906: 184 (ditto); Kato, 1932: 32, text- fig. 37c, 189, pl. 26, fig. 12; Metcalf, 1963: 424 Syn. nov. Drepanopsaltria culta; Breddin, 1901: 28, 113; Jacobi, 1903:10. The following references to Prasia faticina were found to relate to other species: Distant, 1892: 145 (partim: specimen from Kandari is unidentified); Breddin, 1901: 112, 113, 153, pl. 1 fig. 17 (= Lembeja maculosa (Distant, 1883)). The following reference to Prasia culta was found to relate to P. princeps: Lallemand, 1931: 78. Lectotype designation. Prasia culta was described after two male specimens from Patunuang, both stored in the British Museum (Nat. Hist.). One, bearing the | following labels, is designated lectotype: “culta/ Dist.” (handwritten, black); “S. Celebes/ Patu- nuang/ Jan. 1896/ H. Fruhstorfer” (print, black cadre); “Type” (round label, red edged, print); SUN PEN (round label, blue edged; print); “8” (print); “Distant Coll./ 1911—383” (print). Consequently, the other specimen with a syntype label is a paralectotype. Synonymy. Comparison of the female holotype of P. fati- cina and the female P. culta specimens inferred conspecificity, though the holotype of P. faticı- na is of an almost entire olivaceous colour. The original description of P. faticina by Stal (1863) gives as body colour: “Pallidissime subolivaceo- flavescens”, which means that the body had a sort of yellowish colouring. This is an indication that the original coloration has changed. Fur- thermore, the female culta-specimens have a coloration that is much more alike the colora- tion given by Stal (1863). Description. Body of males dark-brown or light orange- brownish, dark coloured specimens with a light pronotum with a dark collar. Females orange- yellowish with olivaceous. Dark male speci- mens ventrally paler. Head and pronotum to- gether 1.02—1.22 X as long as meso- and meta- notum together. Female thorax and head to- gether 0.83—0.94 x as long as abdomen; male 0.73—0.82 x as long. Greatest width of the body at the height of pronotum collar or 3rd ab- dominal segment. Head. — Dark or light in males, light in fe- males. Light coloured males brown between the eyes with a middorsal stripe from margin of -pronotum up to fissure between lateral ocelli. Eyes 0.63—0.8 X as wide as vertex width be- tween eyes. Ocelli raised. Distance between lateral ocelli 1—1.45 X as long as distance be- tween eye and lateral ocellus. Head 1.49—1.85 x as long as vertex width between eyes. Width of head 2.26—2.6 X as wide as width of vertex between eyes. Postclypeus in ventral view strongly laterally compressed, paler than dorsal- ly. Transverse ridges concolorous, sometimes upper four ridges slightly darker coloured. Ros- trum with a black apex reaching hind margin of intermediate coxae. Thorax. — In dark male specimens pronotum collar, lateral margin of pronotum and central fascia chocolate-brown, in lighter specimens this coloration less conspicuous. Females with a dark coloured line on the lateral margin (not reaching the pronotum collar) only. Pronotum collar 1.77—2.02 X as wide as head including eyes. Mesonotum with four pale, irregularly speckled obconical areas, sometimes hardly or not discernable. Lateral parts of mesonotum and parts in front of cruciform elevation sometimes (especially in the females) olivaceous tinged. In dark coloured male specimens poste- rior part of cruciform elevation as well as meta- notum chocolate-brown. Legs. — Concolorous, except fore tibiae and tarsi, which are mostly dark-brown. 3. Tegmina and wings. — Tegmina pale hya- line, extreme base vermillion-red. Costal mem- brane chocolate-brown, in lighter coloured males pale-brown. Venation brownish- or pale- ochraceous, basal area infuscated. Third ulnar area (sometimes shaped as in fig. 20) 0.65—0.83 x as long as Ist ulnar area; 4th ulnar area 0.64— 0.74 X as long as radial area. Apical areas of the tegmen long, 4th, 5th, 7th longest; 3rd apical area 0.58—0.71 X as long as 4th one. Hardly any indication of a corial fold, nor of any rem- nant of a transverse vein extending from the 2nd ulnar area into the 3rd. Wings pale hyaline, col- our of venation as in tegmina, extreme base ver- million-red. Cu, and A, do not fuse. Operculum. — Pale-ochraceous. Slender, pointed apically; meracanthus fairly broad, sharply pointed, reaching beyond proximal part of the operculum. 172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Figs. 8—16. Prasia faticina, 3, Patunuang. 8, pygoter, ventral view (av = anal valves, cdb = caudal dorsal beak, cl = clasper, hplm = hindpart lateral margin, Ip! = lateral pygofer lobe); 9. pygofer, ventrolateral view (aa = aedeagal adjustment, u = uncus); 10, clasper, lateral view; 11, uncus and claspers, ventral view (mpu = median part of uncus); 12, apex of aedeagus, 12a, laterodorsal view, 12b, lateral view; 13, apex of aedeagal appendage process; 14, operculum, ventral view; 15, sternite 8, ventral view; 16, tymbal, lateral view. De Jong: Oriental Prasia 173 Figs. 17—20. Prasia faticina; 17, & Patunuang; 18—20, © Patunuang. 17, edge of sternite 1, ventral view; 18, sternite 7, ventral view; 19, operculum, ventral view; 20, 3rd ulnar area of right tegmen. Abdomen. — Tergites dark- or light-brown, with orange-red hindmargins. In darker speci- mens hindmargins of sternites 3—7 greyish- brown, and tergites with pale lateral spots. Ter- gite 1 trapezoid with sharp proximal and obtuse distal angles. Hindmargin of tergite 1 twice as long as medial length of tergite. Tergite 3 long- est. Sternite 8 apically pointed. Tymbals. — Medium-sized, provided with 8 chocolate-brown long ridges alternating with short orange-brown medial ridges, of which the smallest one is sometimes hardly visible. Genitalia. — Lateral lobes in lateral view swollen and pointed, in ventral view basally swollen, apically slender and pointed. Caudal dorsal beak at a slight angle with dorsal part of the pygofer. Claspers in lateral view very broad and strongly curved apically; hardly elongate. Uncus as in fig. 11, with a small or hardly devel- oped medial protuberance. Aedeagus slender with slender, slightly dentate, short apical lobes. Adjustment of aedeagus situated at half the length of the pygofer. Dorsal aedeagal append- age originating at about 2/3 of aedeagus length and split in two slender, apically pointed, pro- cesses at about 2/3 of its own length, these pro- cesses not reaching apex of aedeagus. 2. Tegmina and wings. — Tegmina subhya- line with an orange-brownish tinge; extreme base pale-red. Costal membrane white. Vena- tion pale-ochraceous, basal area infuscated. Third ulnar area (sometimes shaped as in fig. 20) 0.55—0.74 X as long as Ist ulnar area; 4th ulnar area 0.62—0.66 X as long as radial area. Apical areas as in males, 3rd apical area 0.55—0.69 x as long as 4th one. Hardly any indication of a transverse vein extending from the 2nd ulnar area into the 3rd. Wings pale hyaline, colour of venation as in tegmina, extreme base pale-red. Cu, and A, do not fuse. Operculum. — Pale-ochraceous, rounded in holotype, more of a pointed shape in the other female specimens. Meracanthus as in males. Abdomen. — Orange-brown with sometimes an olivaceous tinge, especially on the weakly ca- rinated medio-dorsal “ridge”. Caudal dorsal beak slender. Ovipositor sheath not reaching beyond caudal dorsal beak. Sternite 7 as in fig. 18.Measurements based upon all specimens available: body length d: 19.6—23.8 mm, x = 22.3, o= 1.727, 9: 24.6—26.2 mm, x= 25.4, o = 0.583; width of pronotum collar 3: 7.2—8.3 mm, x = 7.7, 0 = 0.339, 2: 9— 9.6 mm, x = 9.3, o= 0.286; tegmen length OS Asi nmr V5, U 105 2s 34.9—37.9 mm, x = 36.0, = 1.190. Distribution. — South Sulawesi (Patunuang, Ujung Pandang (= Makassar)) (fig. 7). Material examined. — Indonesia, Sulawesi: Patu- nuang, H. Fruhstorfer, i. 1896, 1 d lectotype of Prasia culta (BM), 1 d paralectotype of Prasia culta (BM), 4 3 2 2 (CNMW), same data but with: 1909—21, Pra- sia sp., 1 2 (BM), same data but with: Prasia Dist; 1910—6, coll. A. Jacobi, 1 & (SMD), same data but with: Prasia culta, 1 & (AMS), same data but with: Drepanopsaltria culta, 1 8 (DEI), same data but with: Drepanopsaltria culta Dist, 1 & (TMB), “Mak” (= Makassar, now Ujung Pandang, blue round label, handwritten), “Celeb/Wallace” (partly print, partly 174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 handwritten), “faticina. Stal” (handwritten), “67/66” (handwritten, blue round label), “syntype” (print, blue edged, round label), 1 2, holotype of Prasia fati- cina (BM). Remarks. P. faticina is closely related to P. breddini n. sp., and differences between these species are only small. Two other very closely related spe- cies are P. tuberculata n. sp. and P. sarasinorum n: sp., which are easier discerned from P. fatici- na. Among the males of P. faticina are very dark and light coloured specimens. Since, how- ever, their genitalia are all alike, and no other structural differences have been found, I have refrained from a taxonomic separation between the two colour forms. Prasia breddini n. sp. (figs. 7, 21—30) Since Prasia breddini resembles P. faticina in almost every respect, a differential description is presented. The species is described after two male specimens. Description of the male. Body of male dark, holotype a little paler than a dark coloured P. faticina specimen. Head and pronotum together 1.1—1.11 X as long as meso- and metanotum together. Thorax and head together 0.77 x as long as abdomen. Greatest width of the body at the height of the 3rd abdominal segment. Head. — Uniformly brown, with a dorsal medial longitudinal pale line on postclypeus. Eyes 0.71—0.79 X as wide as vertex width be- tween eyes. Distance between lateral ocelli 1— 1.25 X as long as distance between lateral ocel- lus and eye. Head 1.69—1.72 X as long as ver- tex width between eyes. Head 2.42—2.58 X as wide as vertex width between eyes. Transverse ridges less distinct than in dark P. faticina speci- mens. Head in lateral view more rounded than in P. faticina. Rostrum reaching intermediate trochanter. Thorax. — Coloration of pronotum interme- diate between dark and lighter coloured P. fati- cina specimens. Pronotum collar 1.91—1.92 x as wide as head including eyes. Mesonotum pale-brown with patches indicating four obcon- ical areas, except for a brown spot in front of cruciform elevation. Ventral surface with thick- er hairs than in P. faticina. Legs. — Shape as in P. faticina, fore tibiae and tarsi only slightly darker than femora. Tegmina and wings. — Third ulnar area 0.63 x as long as 1st one; 4th ulnar area 0.62—0.67 x as long as radial area. 3rd apical area 0.61 X as long as 4th one. Operculum. — As in P. faticina. General ap- pearance only slightly broader. Abdomen. — In holotype paler than the dark coloured P. faticina specimens, in paratype fairly dark. Sternites with broad, orange-brown to red coloured hindmargins. Tymbals. — Eight pairs of alternating ridges, the smallest short ridge is, as may be found in P. faticina, hardly visible. Genitalia. — Pygofer more sturdy than in P. faticina. Claspers elongate and not strongly curved apically; its dark brown apex pointed. Median part of uncus above aedeagus broader than in P. faticina; sometimes provided with a small medial protuberance. Dorsal part of uncus raised high above the lateral margin of the py- gofer. Aedeagus longer and stouter than in P. faticina, its apical lobes longer and less slen- der; dentate. Adjustment of aedeagus at half the length of the pygofer. Pointed processes of dor- sal aedeagal appendage somewhat broader than in P. faticina. Measurements of the d types: body length: 22.3 mm; width of pronotum collar: 7.6—8.2 mm; tegmen length: 29.5 mm. Distribution. — The male holotype has been collected in Ussu, Central Sulawesi, near Malili. The paratype is from Palopo, Central Sulawesi (fig. 7). Types. — Indonesia, Sulawesi: “Celebes/ Us- su/ leg. Dres. Sarasin” (handwritten), “Ussu” (handwritten), “1910/ 6” (partly print, partly handwritten), “coll. A. Jacobi” (print), “Staatl. Museum für/ Tierkunde Dresden” (print), 1 6, holotype of Prasia breddini (SMD); Palopo, Ce- lebes, 1 6 paratype of Prasia breddini (MZB). Etymology. — The species is named after the German hemipterologist Dr. Gustav Breddin, who recognized the separate taxonomic posi- tion of Prasia as defined in his description of Drepanopsaltria. Remarks. Differences between P. breddini and P. fatici- na are mainly found in the genital structures of the two species. As the differences are very slight, the species are probably very closely re- lated. There are several female specimens that may be attributed to P. breddini. The supposed taxo- nomic position of some unidentified females De Jone: Oriental Prasia 175 Figs. 21—28. Prasia breddini, holotype. 21, pygofer, lateral view; 22, clasper, lateral view; 23, pygofer, ventro- lateral view; 24, uncus, ventral view; 25, apex of aedeagus, 25a, laterodorsal view, 25b, lateral view; 26, apex of aedeagal appendage process; 27, edge of sternite 1, ventral view; 28, sternite 8, ventral view. 176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 29 NEE: KL Figs. 29, 30. Prasia breddini, holotype. 29, operculum, ventral view; 30, tymbal, lateral view. will be discussed in the remarks of P. sarasino- rum n. SP. Prasia sarasinorum n. sp. (figs. 7, 31—40, 88) The differential description of P. sarasinorum hereafter is made in comparison with P. fatici- na. The species is described after four male specimens. Description of the male. Body pale-orange-brown. Specimens studied without traces of a dark coloured fascia on the pronotum. Ventrally a little paler than dorsally. Head and pronotum together 1.12—1.21 X as long as meso- and metanotum together. Thorax and head together 0.79 x as long as abdomen. Greatest width of body at the height of either the 3rd abdominal segment or pronotum collar. Head. — Slightly darker between eyes. Eyes 0.78—0.84 X as wide as vertex between eyes. Distance between lateral ocelli 0.91—1.52 X as long as distance between eye and lateral ocellus. Head 1.85—2 X as long as vertex width be- tween eyes. Width of head 2.57—2.68 X as wide as vertex between eyes. Rostrum reaching beyond intermediate coxae. Thorax. — Pronotum unicolorous, some- times a little darker on pronotum collar, espe- cially at the lateral corners. Pronotum collar 1.88—2.01 X as wide as head including eyes. Mesonotum in only one specimen unicolorous, which seems to be the natural coloration, since the other specimens (with odd patches) have been kept in alcohol for some time. Ventral sur- face of thorax with thicker hairs than in P. fati- cina. Legs. — As in P. faticina, but fore tibiae and tarsi less conspicuously darker coloured. Tegmina and wings. — Third ulnar area 0.66—0.71 X as long as Ist one; 4th ulnar area 0.64—0.71 X as long as radial area. Third apical area 0.67—0.69 X as long as 4th one. Operculum. — Broader than in P. faticina; apex distinctly rounded. Abdomen. — Tergites orange-brown, ster- nites only a little paler. Tergites 3—6 only with small red, sternites with broad red hindmargins. Sternite 1 laterally almost straight. Sternite 8 apically less pointed than in P. faticina. Tymbals. — Eight long ridges alternating with short ridges, smallest short one mostly dis- tinctly visible, and even a 9th long ridge sometimes discernable. Genitalia. — Pygofer small compared to P. faticina. Caudal dorsal beak shorter than in P. faticina. Lateral margins of pygofer in be- tween caudal dorsal beak and each of the lateral lobes hardly, though distinctly, degrading. Lateral lobes a little smaller than in P. faticina. Claspers in lateral view more elongate and api- cally dark, hardly curved. Aedeagus in general appearance more sturdy than in P. faticina. Api- cal lobes of aedeagus very short and broad; den- tate. Adjustment of aedeagus situated less than DE Jone: Oriental Prasia 177 32 a À Figs. 31—40. Prasia sarasinorum. 31, pygofer, ventrolateral view, holotype; 32, apex of aedeagal appendage pro- cess, holotype; 33, apex of aedeagus, 33a, laterodorsal view, 33b, lateral view, holotype; 34, pygofer, lateral view, holotype; 35, clasper, lateral view, holotype; 36, uncus, ventral view, holotype; 37, sternite 8, ventral view, holotype; 38, tymbal, lateral view, paratype Mapane; 39, operculum, ventral view, paratype Mapane; 40, edge of sternite 1, ventral view, holotype. 178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 halfway the length of the pygofer. Processes of dorsal aedeagus appendage pointed. Meaurements of the d types: body length: 22.9—23.2 mm (n = 2); width of pronotum col- lar: 7.9—8.3 mm, x = 8.1, 0 = 0.145; tegmen length: 28.5—28.9 mm, x = 28.8, 0 = 0.189. Distribution. — Central Sulawesi (fig. 7). Types. — Indonesia, Sulawesi: “Celebes/ Mapane & Umgebg/ii.95 (Dres. Sarasin)” (handwritten), “Prasia/Distanti/Bred.” (hand- written), “Prasia culta?” (handwritten), „Dis- tant Coll./1911—383.” (print), 1 6, holotype of Prasia sarasınorum (BM); Mapane & surround- ings, Dres. Sarasin, ii. 95, coll. A. Jacobi, 1910— 6, 1 d paratype of Prasia sarasinorum (SMD), | same locality and collector but with: coll. Bred- din, 1 d paratype of Prasia sarasinorum (DEI); Posso (near lake), 11.95, Dres. Sarasin, coll. Breddin, 1 d paratype of Prasia sarasinorum (DEN), Etymology.— This species is named after the Sarasin brothers, in recognition of their contri- butions to our knowledge of the fauna of Sula- wesi by their collecting activities and important publications on the biogeography of the island (e.g. Sarasin & Sarasin, 1901). Remarks. The species is easily recognized by its round- ed opercula, the apex of the aedeagus, and the lacking of dark coloured central fascia on the pronotum. The character last mentioned is shared with P. tuberculata n. sp. The material collected by the Sarasin brothers also contained two females, one from the sur- roundings of Lake Posso and one from the southern headlands of the Takalekadjo Range in Central Sulawesi. These are characterized by a slender tergite 9 and the ovipositor sheath being longer than in any other Prasia female. As there is another female specimen from Tentena, that is different from both specimens collected by the Sarasins, it is impossible to attribute these three females to any species yet, regarding P. sarası- norum, P. senilirata n. sp. and P. breddini in particular, since these three species are recorded from Central Sulawesi. More material may lead to a proper identification of these females. Prasia tuberculata n. sp. (figs. 7, 41—50) The differential description, after the male holotype, is made in comparison with P. fatici- na. Description of the male. Body brown coloured. Ventrally paler than dorsally. Head somewhat darker. Holotype without traces of dark coloured fascia on the pronotum. Head and pronotum together 1.05 x as long as meso- and metanotum together. Tho- rax and head together 0.78 X as long as abdo- men. Greatest width of the body at the height of the 3rd abdominal segment. Head. — Slightly darker between ocelli. Eyes 0.72 X as wide as vertex between eyes. Distance between lateral ocelli 1.2 X distance between lateral ocellus and eye. Head 1.78 X as long as width of vertex between eyes. Width of head 2.43 X width of vertex between eyes. Postcly- peus ventrally pale coloured. Rostrum just reaching intermediate trochanter. Thorax. — Pronotum on the whole unicolo- rous, somewhat darker coloured on the prono- tum collar, especially at the lateral corners. Pro- notum collar 1.86 X as wide as width of head including eyes. Mesonotum, with the four ob- conical areas slightly lighter coloured, concolo- rous. Cruciform elevation slightly darker col- oured. Legs. — As in P. faticina, the fore tibiae and tarsi only being slightly darker than the remain- der. Tegmina and wings. — Costal membrane not as dark as in P. faticina. Third ulnar area (shaped as in fig. 20) 0.81 X as long as 1st one; 4th ulnar area 0.66 X as long as radial area. Api- cal areas 4, 5, 6 and 7 longest. Third apical area 0.66 X as long as 4th one. Operculum. — As in P. faticina, meracanthus more slender. Abdomen. — Tergites brown, sternites only a little paler. Tergites 3—6 only with small red, sternites with broad pale-red hindmargins. Ster- nite 8 far less pointed. Tymbal. — Seven long ridges alternating with an equal number of short ridges. Genitalia. — Pygofer small compared to P. faticina. Caudal dorsal beak mutilated. Lateral margins of pygofer hardly, though dis- tinctly, degrading. Lateral lobes a little smaller, in ventral view more pointed. Claspers in lateral view apically slender, but with a distinct swell- ing dorsally. Median uncus part small, without protuberance. Apex of aedeagus longer than in P. faticina; aedeagus slender. Adjustment of ae- deagus situated slightly lower in comparison with P. faticina. Processes of dorsal aedeagal appendage very slender and pointed apically. Measurements of the holotype: body length: DE Jone: Oriental Prasia 179 Figs. 41—48. Prasia tuberculata, holotype. 41, uncus, ventral view; 42, pygofer, lateral view; 43, clasper, lateral view; 44, pygofer, ventrolateral view; 45, apex of aedeagal appendage process; 46, apex of aedeagus, 46a, latero- dorsal view, 46b, lateral view; 47, operculum, ventral view; 48, sternite 8, ventral view. 180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Figs. 49, 50. Prasia tuberculata, holotype. 49, tymbal, lateral view; 50, edge of sternite 1, ventral view. 22.8 mm; width of pronotum collar: 7.6 mm; tegmen length: 27.4 mm. Distribution. — The holotype is from Tom- bugu (= Tombuko) in East Sulawesi (fig. 7). Type. — Indonesia, Sulawesi: “Ost-Cele- bes/Tombugu/H. Kühn 1885” (print, black cadre), 1 6 (MNP). Etymology. — The species is named after its swelling on the clasper, when looked at lateral- ly. “Tuberculata” is Latin for swollen. Remarks. At first sight this species is easily mixed up with P. sarasinorum, because of the lacking of coloration of the central fascia on the prono- tum. Its genitalia, however, are very different from the species mentioned, and its opercula are pointed, whilst in P. sarasinorum they are rounded apically. The apex of the aedeagus seems a sort of combination between P. faticina and P. sarasinorum. The number of long ridges is the same as in P. nigropercula. Prasia nigropercula n. sp. (figs. 6, 7, 51—59, 89) The description is made in comparison with P. faticina and based upon the single male speci- men from Muna Island, situated near the south- eastern arm of Sulawesi. Description of the male. Body fairly dark coloured, especially head, pronotum collar, central fascia and obconical spots on mesonotum. Head and pronotum to- gether 1.34 X as long as meso- and metanotum together. Head and thorax together 0.78 X as long as abdomen. Greatest width of body at the height of pronotum collar. Head. — Dark-brown, ocelli on a black un- derground. Head (a little damaged) much por- rect. Eyes 0.67 X as wide as vertex width be- tween eyes. Distance between lateral ocelli 1.1 x distance between eye and lateral ocellus. Head 1.82 X as long as width of vertex between eyes. Head 2.34 X as wide as vertex between eyes. Postclypeus in ventral view darkening dis- tad. Rostrum reaching intermediate coxae. Thorax. — Pronotum collar, lateral margin of pronotum, central fascia and some spots on the pronotum dark-brown coloured. Pronotum col- lar more or less pointed at the lateral corners, 1.96 X as wide as head including eyes. Mesono- tum with sıx dark-brown coloured areas at proximal margin, two paramedian ones half as long as more lateral ones, which join broadly in front of cruciform elevation. Cruciform eleva- tion dark-brown for its greater part. Metano- tum dark-brown. Ventrally pale ochraceous. Legs. — As in P. faticina. Tegmina and wings. — Venation and costal membrane dark-brown. Third ulnar area 0.7 X as long as 1st one; 4th ulnar area 0.67 X as long as radial area. Third apical area 0.66 X as long as 4th one. Traces of coloration of transverse vein present. Operculum. — Dark, nearly black. Smaller than in P. faticina; acutely pointed. Meracan- thus dark coloured, size as in P. faticina. DE Jong: Oriental Prasia 181 Figs. 51—56. Prasia nigropercula, holotype. 51, pygofer, lateral view; 52, apex of aedeagal appendage process; 53, pygofer, ventrolateral view: 54, uncus, ventral view: 55, clasper, lateral view; 56, edge of sternite 1, ventral view. Abdomen. — Dark-brown coloured with red hindmargins, along tergites as well as sternites. Tergite 1 for its greater part covered by metano- tum. Sternite 1 apically somewhat smaller and laterally more concave than in P. faticina; medi- an protuberance more conspicuous. Sternite 8 hardly pointed apically. Tymbals. — Seven long ridges alternating with an equal number of short medial ridges. Genitalia. — Caudal dorsal beak almost in a straight line with dorsal part of pygofer. Lateral lobes smaller than in P. faticina. Median uncus part above aedeagus broad. Claspers slender, somewhat elongate, hardly curved and apically 182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Figs. 57—59. Prasia nigropercula, holotype. 57, tymbal, lateral view; 58, sternite 8, ventral view; 59, operculum, ventral view. rounded. Aedeagus extremely slender (apex mutilated). Adjustment of aedeagus situated less than halfway the pygofer. Processes of dorsal aedeagal appendage apically rounded. Measurements of the holotype: body length: 23.4 mm; width of pronotum collar: 8.4 mm; tegmen length: 28.4 mm. Distribution. — Muna (Raha) (fig. 7). Type. — Indonesia, Muna: “Raha/Moena/Ile Celebes” (handwritten), 1 d (BIN). Etymology. — The species is named after its nearly black coloured operculum, that contrasts with the pale underside of the body. Remarks. Beside its conspicuous coloration, the species is very distinct within the genus because of some characteristic details in the genitalia struc- tures, viz. the shape of the claspers and the ad- justment of the aedeagus, as well as the apically rounded processes of the aedeagal appendage. The shape of the clasper, though, is reminiscent of that of P. sarasinorum. A female from Kandari (= Kendari?), which has probably lost the natural coloration, has about the pronotum collar shape of P. nigroper- cula. Since the material is too scanty, this speci- men is still regarded as unidentified. Prasia princeps Distant, 1888 (figs. 7, 60—75, 90, 91) Prasia princeps Distant, 1888: 325; Distant, 1892: xiv, 145, pl. 13, figs. 14, 14a, b; Jacobi, 1903: 12; Dis- tant, 1906: 184 (equals Drepanopsaltria (?) prin- ceps Bredd.); Kato, 1932: 189; Metcalt, 1963: 425. Drepanopsaltria (?) princeps; Breddin, 1901: 28, 113. Drepanopsaltria princeps; Jacobi, 1903: 10. Description. Body olivaceous-green to green, sometimes with a brownish tinge. Head darker coloured. Body ventrally somewhat paler than dorsally. Head and pronotum together 0.96—1.22 X as long as meso- and metanotum together. Female thorax and head together 0.81—1.02 X as long as abdomen; male 0.69—0.75 X as long. Great- est width of the body at the height of the 3rd abdominal segment. Head. — Dorsally olivaceous with brown be- tween eyes. Eyes large, 0.67—0.88 X as wide as vertex width between eyes. Ocelli raised. Dis- tance between lateral ocelli 0.86—1.53 x dis- tance between eye and lateral ocellus. Head 1.63—1.98 X as long as width of vertex between eyes. Head 2.35—2.76 X as wide as vertex be- tween eyes. Postclypeus in ventral view strong- ly laterally compressed, olivaceous- to light- brown coloured. Transverse ridges weak, con- colorous. Rostrum with black apex reaching in- termediate coxae. Thorax. — Central fascia on pronotum collar obsolete; pronotum collar broadly rounded, 1.84-2.11 X as wide as width of head including eyes. Fissures on pronotum undeep. A lateral brown line running from each eye backwards, almost reaching latero-proximal corner of pro- notum collar. Mesonotum with four speckled obconical areas at proximal margin; the para- DE Jone: Oriental Prasia 183 Figs. 60—67. Prasia princeps, 3. 60, 61, clasper, lateral view, 60, Menado, 61, Toli-Toli; 62, 63, pygofer, ventro- lateral (62) and lateral (63) view, Tanggarie-Menado; 64, 65, apex of aedeagal appendage process, 64, Toli-Toli, 65, Menado; 66, apex of aedeagus, 66a, laterodorsal view, 66b, lateral view, Toli-Toli; 67, apex of aedeagus, 67a, laterodorsal view, 67b, lateral view, Menado. 184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Figs. 68—70. Prasia princeps: 68, 69, 3; 70, ©. 68, uncus, ventral view, Menado; 69, edge of sternite 1, ventral. | view, Menado; 70, 3rd ulnar area of right tegmen, Toli-Toli. median areas being half as long as the lateral ones, whose length is about 3/4 of the disk. Cruciform elevation fairly flattened. Metano- tum just extending from below mesonotum. Legs. — Concolorous, except fore tibiae and tarsi, which are brownish. 3. Tegmina and wings. — Tegmina very pale-ochraceous or olivaceous, subhyaline. Costal membrane olivaceous. Extreme base red. Apical areas of tegmen long, 4th, 5th and 6th longest, 1st, 3rd and 7th shorter, 2nd and 8th shortest. Third apical area 0.58—0.69 X as long as 4th one. Third ulnar area (sometimes shaped as in fig. 70) 0.7—0.87 X as long as 1st one; 4th ulnar area 0.63—0.76 X as long as radial area. Hardly any indication of a corial fold, nor of any remnant of a transverse vein extending from the 2nd ulnar area into the 3rd. Wings pale hyaline, venation ochraceous, ex- treme base red. Cu, and A, veins fused just be- fore the wing border. Operculum. — Large, sickle-shaped, broad and pointed, just reaching or reaching just be- yond sternite 2, concolorous. Meracanthus short and slender, not reaching beyond proxi- mal part of operculum. Abdomen. — Sometimes irregularly speckled with light patches, fresh specimens unicolorous. Tergite 1 trapezoid with sharp proximal and ob- tuse distal angles. Hindmargins of tergites orange-brown coloured. Distal edge of sternite 1 convex with a weak median bulb. Sternite 8 apically slightly to hardly pointed. Tymbals. — Medium-sized and consisting of 9 (sometimes 10 and one male from Toli-Toli 8) long ridges alternating with short medial brown ridges in a very regular pattern. Genitalia. — Lateral lobes of the pygofer in lateral view swollen and pointed, in ventral view swollen, apically slender and pointed. Caudal dorsal beak sometimes at an angle of at most 40° with dorsal part of pygofer. Claspers in lateral view broad, sometimes more narrowly shaped, somewhat elongate, apically curved and point- ed; in dorsal view median part slender and api- cally swollen. Uncus consisting of two lateral, slightly swollen parts and mostly of a medial protuberance. Greater part of the aedeagus slen- der. Dorsal aedeagal appendage originating at about 2/3 of the aedeagal length, and split at about 2/3 of its own length in two slender pro- cesses, which are apically rounded or pointed. These processes not reaching apex of aedeagus. The rounded ones mostly slightly constricted subapically. Apex of aedeagus consisting of two lateral slender, or somewhat broader lobes, that are dentate apically. Adjustment of aedeagus situated halfway the pygofer. 2. Tegmina and wings. — Tegmina pale- ochraceous or olivaceous, subhyaline. Costal membrane white. Extreme base red. Apical areas as in males. Third apical area 0.61—0.71 x as long as 4th one. Third ulnar area (sometimes shaped as in fig. 70) 0.59—0.76 X as long as Ist one; 4th ulnar area 0.67—0.77 X as long as ra- dial area. Hardly any indication of the corial De Jong: Oriental Prasia 185 Figs. 71—75. Prasia princeps; 71—73, 3; 74, 75, 2. 71, sternite 8, ventral view, Menado; 72, operculum, ventral view, Minahassa; 73, tymbal, lateral view, Menado; 74, sternite 7, ventral view, Menado; 75, operculum, ventral view, Woloan-Menado. fold, nor of any remnant of a transverse vein ex- tending from the 2nd ulnar area into the 3rd. Wings pale-hyaline, venation ochraceous, ex- treme base pink. Cu, and A, fused in nearly all specimens (including the type!) just before the wing border. Operculum. — Short, more or less broadly rounded; distal part sometimes shorter than the basal part. Meracanthus reaching just beyond posterior margin of operculum. Abdomen. — Olivaceous, greenish, some- times slightly brownish tinged, fresh specimens green. Broad tergites, weakly carinate medially. Caudal dorsal beak slender. Ovipositor sheath just reaching apex of caudal dorsal beak. Ster- nite 7 as in fig. 74. Measurements based upon all specimens available: body length d: 24.6—28.9 mm, x = 27:3, 01.016, Pre 25,9 39.2, mm, x = 28.2, 6 = 1.394; width of pronotum collar ds 8.0—95 mm, x= 90, o= 0.392, ©: 9.3—11.3 mm, x= 10.2, o = 0.583; tegmen length d: 32—36.7 mm, x = 34.5, o = 1.223, ? : 39.1—41.9 mm, x = 40.6, 0 = 0.819. Distribution. — North Sulawesi (fig. 7). 186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Material examined.— Indonesia, Sulawesi: Dumo- ga-Bone N.P., Sulawesi Utara, Project Wallace 1985, st. 4, lowland rainforest near base camp, 29.i— 2.1.1985, at M.V. light, J. P. Duffels & J. D. Hollo- way collectors, 1 2 (MZB), same locality and collec- tors but with, st. 7, lowland rainforest at 120 m from st. 4, 12.1.1985, at M.V. light, 1 2 (ZMA), 1 2 (MZB), same locality and collectors but with, st. 9, subcamp 1, 4—8.11.1985, at light, 3 d (ZMA), 4 & (MZB); Menado, van Braeckel, 1 d 1 2 (BIN); Mena- do, van Braeckel, Prasia culta Dist., det. Lallemand 1930, 3 d (BIN); Minahassa, “Minahassa/Celebes” (handwritten), “Syntype” (round label, blue edged, print), “Distant Coll./1911—383” (print), 1 2, holo- type of Prasia princeps (BM); Minahassa, Prasia prin- ceps Dist., coll. Dr. D. MacGillavry, 1 2 (ZMA); Minahassa, V. d. Bergh, Prasia faticina Stal, coll. Dr. D. MacGillavry, 1 d (ZMA); Tanggarie-Menado, Van Braeckel, Prasia culta Distant, det. Lallemand 1930, 1 d (BIN); Toli-Toli, Nord-Celebes, Nov.- Dez. 1895, H. Fruhstorfer, 1 d (CNMW), same data but with, 1909—21, 1 © (BM), same data but with Prasia faticina Dist. 145.vii.14 (= Distant, 1892: 145, pl. 7, fig. 14), 1 6 (MHNG), same data but with, Pra- sia princeps dist. 145.x11.14 (= Distant, 1892: 145, pl. 13, fig. 14), 1 2 (MHNG); Tondano-Menado, Van Braeckel, Prasia culta Distant, det. Lallemand, 1 3 (BIN); Woloan-Menado, 3 d 1 2 (BIN). Specimens without further precision of the locality: Celebes, 1 4 BIN; India Archipel, 1 2 CNMW. Remarks. At first there has been some hesitation whether or not to attribute a separate taxonom- ic position to the specimens from Toli-Toli. Whilst the specimens from Menado display a broad clasper, a very slender apex of the aedea- gus and apically rounded processes of the ae- deagal appendage (with the subapical constric- tion), the Toli-Toli specimens possess a narrow clasper, a broader shaped apex of the aedeagus and pointed processes of the aedeagal append- age. Recent collecting in the Dumoga-Bone N.P. in Sulawesi Utara (during the Project Wallace Expedition) by Dr. Duffels and Dr. Holloway (Commonwealth Institute of Entomology, Lon- don) provided new material that displayed the Menado-type of genitalia as well as a mixture of the Menado- and the Toli-Toli-type. More material from Toli-Toli must prove the stability of its combination of genital characters in order to reconsider a separate taxonomic position. Prasia senilirata n. sp. (figs. 7, 76—85) As Prasia senilirata resembles P. princeps in general appearance, mainly because of its size and the olivaceous-green colouring of the body, a differential description is presented based upon the male holotype. Due to its large size the magnification ratio’s of all drawings of P. semili- rata are 0.75 X those used for the other species. Description of the male. Large species. Body colour (fresh) oliva- ceous-green. Head a little darker coloured. Body ventrally somewhat paler than dorsally.’ Head and pronotum 1.09 x as long as meso- and metanotum together. Thorax and head to- gether 0.76 X as long as abdomen. Greatest width of the body at the height of the 3rd ab- dominal segment. Head. — Dark-green with brown between the eyes. Eyes 0.86 X as wide as vertex width between eyes. Distance between lateral ocelli 1.57 x distance between lateral ocellus and eye. Head 2.07 X as long as and 2.72 X as wide as width of vertex between eyes. Postclypeus dark-coloured. Thorax. — Pronotum collar 1.94 X as wide as head including eyes. Dark line running from the eye hindwards very short. Legs. — As in P. princeps. Tegmina and wings. — Coloration as in P. princeps (left tegmen of holotype has 7 apical areas). Third apical area 0.7 X as long as the 4th one. Third ulnar area 0.83 X as long as the Ist one; 4th ulnar area 0.71 X as long as the radial area. Coloration of wings as in P. princeps. Cu, and A, veins do not fuse. Operculum. — Relatively not as large and broad as in P. princeps; concolorous. Abdomen. — Coloration as in olivaceous- green P. princeps specimens. Distal edge of ster- nite 1 smaller and far less bulbed than in P. prin- ceps. Sternite 8 weakly pointed. Tymbals. — Relatively small, consisting of 6 pair of long ridges alternating with short ridges. Upper half of the tymbal shaded with black above the small ridges. Genitalia. — Lateral lobes less swollen than in P. princeps. Caudal dorsal beak at a very small angle with dorsal part of the pygofer. Clasper more elongate than in P. princeps. Me- dian uncus part very slender, without protuber- ance. Aedeagus slender. Processes of dorsal ae- deagal appendage apically pointed. Apex of ae- deagus with two broadened flaps, that have a slight curvature. Measurements of the holotype: bedy length: 29.7 mm; width of pronotum collar: 9.9 mm; tegmen length: 36.2 mm. DE Jon: Oriental Prasia 187 Figs. 76—82. Prasia senilirata, holotype. 76, pygofer, ventrolateral view; 77, apex of aedeagus, 77a, laterodorsal view, 77b, lateral view; 78, apex of aedeagal appendage process; 79, pygofer, lateral view; 80, uncus, ventral view; 81 clasper, lateral view; 82, edge of sternite 1, ventral view. 188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 Figs. 83—85. Prasia senilirata, holotype. 83, operculum, ventral view; 84, sternite 8, ventral view; 85, tymbal, lateral view. Distribution. — The holotype is from Central Sulawesi (Lore Lindu National Park) (fig. 7). Type. — Indonesia, Sulawesi: “Stat. 44/Low- land/rainforest/ML-light” (print), “Toke Pan- gana/700 m/4 km NE Gimpu/16.111.1985/J. P. & M. J. Duffels” (print), “Indonesia/Sulawesi Tengah/Lore Lindu N.P.” (print), 1 d (ZMA). Etymology. — “Senilirata” is derived from the Latin words “seni”, meaning “each time six of” and “lirata”, meaning “ridge made by a plough”. The combination stands for the fact that the species has six long ridges on the tym- bal, which is characteristic, not only within the genus, but also within the Oriental Prasiini as a whole. Remarks. The species is very distinct, not only for its low number of alternating ridges on the tymbal, but also for its peculiar shaped apex of the ae- deagus. As for a possible female representative of the species the reader is referred to the re- marks at the end of the description of P. sarası- norum. REFERENCES Ashton, H., 1912. Some new Australian Cicadidae. — Proc. R. Soc. Vict., (N.S.) 24: 221—229, pls. 49— Sil. Boulard, M., 1975. Les Plautillidae, Famille nouvelle d’Homopteres Cicadoidea. — Annali Mus. civ. Stor. nat. Giacomo Doria 80: 313—318, figs. 1— Da Breddin, G., 1901. Die Hemipteren von Celebes. Ein Beitrag zur Faunistik der Insel. — Abh. natur- forsch. Ges. Halle 24: 1—213, pl. 1. Distant, W. L., 1883. Contributions to a proposed Monograph of the Homopterous Family Cicadi- . dae. — Part 1. — Proc. zool. Soc. Lond. 1883: 187—194, pl. 25. Distant, W. L., 1888. Descriptions of new species of Oriental Cicadidae. — Ann. Mag. nat. Hist. (6) 2: 323—325. Distant, W. L., 1892. A monograph of Oriental Cica- didae, 5—7: i—xiv, 97—158, pls. 10—15. — Indi- an Museum, London. Distant, W. L., 1898. Description of two new species of Oriental Cicadidae. — Ann. Mag. nat. Hist. (7) 1972 Distant, W. L., 1905. Rhynchotal notes. — XXXV. — Ann. Mag. nat. Hist. (7) 16: 265—280. Distant, W. L., 1906. A synonymic catalogue of Ho- moptera. Part 1 Cicadidae: 1—207. — British Mu- seum, London. Distant, W. L., 1909. New Malayan Rhynchota. — Trans. R. ent. Soc. Lond., 1909: 385—396, pl. 10. Duffels, J. P., 1977. A revision of the genus Diceropy- ga Stal, 1870 (Homoptera, Cicadidae). — Mono- grafieén Ned. ent. Veren. 8: 1—227, figs. 1—265. Duffels, J. P., 1983a. Taxonomy, phylogeny and bi- ogeography of the genus Cosmopsaltria Stal with remarks on the historic biogeography of the sub- tribe Cosmopsaltriaria (Homoptera, Cicadidae). — Pacif. Insects Monogr. 39: 1—127, figs. 1— 135, pls. 1—6. Duffels, J. P., 1983b. Distribution Patterns of Orien- tal Cicadoidea (Homoptera) East of Wallace’s Line and Plate Tectonics. In: Thornton, I.W.B. (ed.), Symposium on Distribution and Evolution of Pacific Insects, Geojournal 7 (6): 491—498. Duffels, J. P. & P. A. van der Laan, 1985. Catalogue of the Cicadoidea (Homoptera, Auchenorhyncha) 1956—1980. — Series entomologica 33: i—xvi, 1—414. Holloway, J. D., 1979. A survey of the Lepidoptera, biogeography and ecology of New Caledonia. — Series entomologica 15: 1—xu, 1—588, figs. 1— 153, pls. 1—87. DE Jone: Oriental Prasia 189 Figs. 86—88. General facies. 86, Prasia‘faticina 3, paratype P. culta; 87, Prasia faticina ©, holotype; 88, Prasia sarasinorum 8, paratype Mapane surroundings. Horvath, G., 1913. Etude morphologique sur la con- nidae). — Bijdr. Dierk. 52 (2): 175—185, figs. 1— struction de l’élytre des Cicadas. — Int. Congr. 23% Ent. 2: 422—432, figs. 7, 8. Jong, M. R. de & J. P. Duffels, 1981. The identity, Jacobi, A., 1903. Uber Singcikaden von Ost-Neugui- distribution and synonymy of Lembeja papuensis nea. — Sber. Ges. naturf. Freunde Berl. 1903: Distant, 1897 (Homoptera, Tibicinidae). — Bull. 10—15, figs. 1—5. zool. Mus. Univ. Amsterdam 8 (7): 53—62, figs. Jong, M. R. de, 1982. The Australian species of the 1—12. genus Lembeja Distant, 1892 (Homoptera, Tibici- Karsch, F. A. F., 1890a. Beiträge zur Kentniss der 190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 2, 1985 91 Figs. 89—91. General facies. 89, Prasia nigropercula 3, holotype; 90, Prasia princeps 3, Minahassa; 91, Prasia princeps 2, holotype. DE Jong: Oriental Prasia 191 Singeikaden Afrika’s und Madagascar’s. — Berl. ent. Z. 35: 85—130, pls. 3, 4. Karsch, F. A. F., 1890b. Ueber die Singcicaden-gat- tung Perissoneura Distant. — Ent. Nachr. Berlin 16: 190—192. Kato, M., 1932. Monograph of Cicadidae: 1—450, pls. 1—32, text-figs. 1—122. Kato M., 1956. The biology of Cicadas: 319 p., 146 figs., 46 pl. — Iwasaki Shoten, Tokyo. Lallemand, V., 1931. Hemiptera-Homoptera. Resul- tats scientifiques du voyage aux Indes Orientales Néerlandaises de LL. AA. RR. le Prince et la Princesse Léopold de Belgique. — Mém. Mus. r. Hist. nat. Belg. 4: 71—85. Metcalf, Z. P., 1963. General catalogue of the Ho- moptera, VIII. Part 2. Tibicinidae: i—iv, 1—492. — North Carolina State College, Raleigh, M. C. Myers, J. G., 1928. The morphology of the Cicadidae (Homoptera). — Proc. zool. Soc. Lond. 1928: 365—472, figs. 1—75. Myers, J. G., 1929. Insect Singers. A Natural History of the Cicadas: 1—xix, 1—304, figs. 1—116, pls. 1—7. — George Routledge and Sons, Ltd, Lon- don. Sarasin, P. & F. Sarasin, 1901. Materialen zur Natur- geschichte der Insel Celebes. Dritter Band: Ueber die geologische Geschichte der Insel Celebes auf Grund der Tierverbreitung (mit 15 Tafeln in Lithographie): i—vi, 1—169, text-figs. 1—5, pl. 1—15 figs. 1—45. — W. Kreidel’s Verlag, Wies- baden. Schmidt, E., 1925. Zwei neue Singcikaden von der In- sel Sumba. — Societas ent. 40: 42, 43. Stal, C., 1862. Synonymiska och systematiska anteck- ningar Ofver Hemiptera. — Ofvers. K. Vetensk. — Akad. Forh. 19: 479—504. Stal, C., 1863. Hemipterorum exoticorum generum et specierum novarum descriptiones. — Trans. R. ent. Soc. Lond. (3) 1:571—603. Stal, C., 1870. Hemiptera insularum Philippinarum. — Bidrag ull Philippenska öarnes Hemipter-fau- na. — Ofvers. K. Vetensk.-Akad. Förh. Stockh. 27 (7): 607—776, pls. 7—9. Walker, F., 1858. Supplement. List of the specimens of Homopterous insects in the collection of the British Museum, 1858: 1—307. Walker, F., 1868. Catalogue of the Homopterous in- sects collected in the Indian Archipelago by Mr. A. R. Wallace, with descriptions of new species. — Proc. Linn. Soc. Lond. (Zool.) 10: 82—193, plese Author’s address: Institute of Taxonomic Zoology, P.O. Box 2015, 1000 HC Amsterdam, The Netherlands. # Hay Ty er ENT fr ar GP Ls: AE EN 7 a 0 la RR RAS a u u = È ri È A od J ta DEEL 128 APLEVERING 3 1985 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DEINE DEREANDSE ENTOMOLOGISCHE VERENIGING INHOUD J. C. Roskam. — Evolutionary patterns in gall midge — host plant associations (Diptera, cecidomyiidae), pp. 193—213, figs. 1—3. Tijdschrift voor Entomologie, deel 128, afl. 3 Gepubliceerd 20-XII-1985 Re fs ie Ba) Car ‘ei ur: at Fa EVOLUTIONARY PATTERNS IN GALL MIDGE — HOST PLANT ASSOCIATIONS (DIPTERA, CECIDOMYIIDAE) by J.C. ROSKAM Division of Population Biology, University of Leiden, The Netherlands ABSTRACT Host plant associations of mainly West Palaearctic gall midges have been analyzed to ex- plain some of the radiation of this highly specialized group of endophytophagous insects. Gall midges behave according to some predictions formulated for phytophagous insects in general: woody host plants accumulate more gall midge species than herbaceous ones. In other aspects gall midges seem to be different: taxonomical affinity of host plants might be more important to explain radiation in gall midges than it is for other groups of plant feed- ers, especially external feeders. Furthermore, gall midges deserve particular attention be- cause the gall inducing feeding mode in this group might be a result of polyphyletical devel- opment. Specialization on host plant organs has been analyzed to support this assumption. Finally, various evolutionary processes allied with insect—host plant interactions have been analyzed for gall midges. Sequential evolution could be demonstrated in this group and some examples of apparent parallel cladogenesis, each dealing with a different rank of host plant taxonomy, are treated. INTRODUCTION According to recent estimates, about 792,000 species of insects have been described, of which 46% feed upon plants (Southwood, 1978; Price, 1977). Important pioneer work on insect—host plant interactions has been done by Verschaffelt (1910), Dethier (1954) and Fraenkel (1959). The enormous expansion of literature on this subject began with the classic papers by Ehrlich & Rav- en (1964) on co-evolution, MacArthur & Wil- son’s (1967) theory of island biogeography and Janzen’s (1968) application of the latter theory to insect-host plant interactions. Recent books by Crawley (1983) and Strong et al. (1984) offer a thorough introduction to the literature on this subject. Plant chemistry and, because related plant taxa often share similar compounds, plant taxonomy played an important role in earlier studies. Gradually more host plant traits be- came involved to explain accumulations of in- sect species on host plant taxa. Fowler & Law- ton (1982), for example, used no less than nine variables, a potpourri of characteristics of host plants, phytophages and natural enemies of phytophages in a multiple regression calculation to explain the species richness of leafminers on British Umbelliferae. In the latter study, host plant taxonomy is not even a significant factor 193 anymore: 61% of the variation is explained by habitat diversity and leaf form of the host plants. In another study, however, about leaf- miners on British trees, 36% of the variation was caused by taxonomical diversity alone, geo- graphic range being the second trait in impor- tance (Godfray, 1982). When the literature is subdivided according to the different guilds of phytophagous insects, it is remarkable that papers dealing with exter- nal plant feeders (chewing and sucking insects) are abundant, whereas references on endophy- tophages, such as miners and gall insects are scarce. Nevertheless it is obvious that not only among external plant feeders, but also among endophytophages there are many species with an important impact on host plant development and seed production, in natural situations (e.g., Harnett & Abrahamson, 1979), as well as in pest control (e.g., Bess & Haramoto, 1959) and in agriculture (e.g., Skuhravy et al., 1983). Fur- thermore, in important aspects endophyto- phages differ basically from external plant feed- ers and deserve therefore special attention. Gall insects in particular not only depend on plants for nutriment, but also for shelter, which is con- structed by manipulating the defense reactions of the host plants. This very precise tuning of 194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 the insect’s needs to the plant’s potentials may explain why polyphagous gall insects (species attacking host plants belonging to different fam- ilies) are extremely scarce, whereas monopha- gous species are abundant. For this reason eco- logical opportunists (species shifting to new host plants which are in close proximity to, rather than taxonomically related with, the orig- inal ones) may be rare too among this group. A further consequence of the gall inducing feeding mode is that colonization of new resources by these insects, but also by miners, is a substan- tially slower process (Strong et al., 1984). Therefore, long term processes, playing in “evolutionary time”, rather than short term processes, in “ecological” time, seem to be more apparent in interactions between host plants and gall midges. The present study deals with host plant rela- tions of mainly West Palaearctic gall midges. Compared with other groups of endophyto- phages, gall midges have some advantages. As a group, they have a broad host plant spectrum, whereas cynipids, another main group of gall insects, are abundant on Fagaceae and Rosaceae only. Agromyzids are also an important group of endophytophages, but are restricted to par- ticular plant organs, mainly leaves. Until now, it has been impossible to analyze host plant relations of gall midges on a world basis since the detailed knowledge needed for such a study is only available for some parts of the temperate regions. Therefore this study is restricted to data presented by Buhr (1964— 1965) and Skuhrava (in press) for the West Pal- aearctic, extended in some cases, where infor- mation from the Nearctic was necessary, to Felt (1940) and Gagné (1969, 1981). Whether the re- sults will hold for other climatic areas must be considered in future. In the next section some main characteristics will be given of the ecology and taxonomy of gall midges. Which life history traits of gall midges are important in connection to host plant relations and change of host plants? How are the Cecidomyiidae, the family gall midges belong to, subdivided and which group(s) gave rise to gall inducers? Gall midges may be unique among gall insects, because arguments exist for a polyphyletic transition of Cecidomyiidae to the gall inducing feeding mode (Mamaev, 1968). A comparison will be made for subgroups of gall midges to investigate whether specialization to different host plant organs (vegetative or gen- erative) may contribute evidence for such a polyphyletic shift. Subsequently, our attention will be focussed on host plant diversity. Because an analysis of evolutionary aspects is our main goal, only tax- onomical and some structural diversity of host plants will be considered to explain radiation of gall midges. Taxonomically diverse plant fami- lies, including many species, are supposed to support more midge species than less diverse families, because there is more scope for adap- tive radiation among phytophages in diverse taxa (Crawley, 1983). Also we will contrast woody against herbaceous host plants, because the first live longer and may be structurally more diverse, and are therefore a more predict- able resource offering again more opportunities for adaptive radiation (Lawton, 1983). Other traits of host plants, such as geographical range, . local abundance and habitat diversity, important ecological variables indeed, must be omitted be- cause accurate scoring is only possible for some local areas, but not for the West Palaearctic as a whole. Finally, knowing something about interac- tions between structural and taxonomical traits of host plants and gall midge diversity, ques- tions rise about the consequences for the evolu- tion of these phytophages. Some interactions re- sulted in the occurrence of related midge species on related groups of host plants. But how abun- dant are apparent parallel patterns in the clado- genies of gall midges and host plants and to what extent did they evolve? Are examples of parallel cladogenesis the result of plant — gall midge interaction, or were the host plants changed under influence of other selection fac- tors and did the gall midges follow these changes? We will draw up examples of parallel cladogenesis and discuss the processes. LIFE HISTORY PATTERNS Knowledge of life history patterns is essential for evolutionary studies because each mode of speciation needs particular prerequisites of the involved organisms. Speciation processes of gall midges, which are relevant here, are those in which host plants are involved. Modes of sym- patric speciation might exist when host plant shifts occur and assortative mating can be dem- onstrated. Partners are preferred which share the same food plant, or a highly similar food resource, during the larval phase. Therefore, mating site, oviposition site and site of larval de- Roskam: Gall midge — host plant associations 195 velopment have to be coupled by localization on the same host. A transfer to a new host re- sults then not only in a new resource, but chan- nels the gene flow by separating mating and ovi- position sites of original and shifted populations (Bush, 1975; Zwolfer & Bush, 1984). On the other hand, modes of allopatric speciation may result from co-evolution, as a reciprocal process between host plants and phytophages or, when the impact of phytophages on host plant changes is doubtful or absent, sequential evolu- tion (Jermy, 1976). Also co-evolution and se- quential evolution require a highly coupled niche structure, but host plant shifts are absent. Therefore, cladogenesis of both groups of organisms is characterized by corresponding di- chotomies (Regenfuss, 1978). In order to inves- tigate which modes of speciation may occur in gall midges, relevant phases of the gall midge life history are analyzed. Gall midges!) alternate a sedentary phase, en- capsulated in a gall, with a free-living adult phase, in which dispersion is possible. The free- living phase starts with the emergence of the adults. Males usually emerge some hours earlier than females and periods of activity are species specific (Coutin & Harris, 1968; Jones et al., 1983; Skuhravy & Skuhrava, 1982). After a short period of rest males start swarming in search of females; usually they hover in groups in close proximity to galls where emerging fe- males are expected. Males may mate several times (Van Vreden & Arifin, 1977). Females, like males, rest for a while after emergence. During this period the ovipositor is extended in a calling position, emitting sex pheromones (McKay & Hatchett, 1984). Attracted males co- pulate immediately, without any courtship be- haviour. Females mate once, after mating they retract the ovipositor and are not receptive any more. The mating, or “rendez-vous” site depends on the site where pupation occurs and, conse- quently, the female emerges. Before pupation, mature larvae either drop onto the soil or re- main in the gall. Galls, in their turn, either may be shed from the host plant or may remain con- nected with it. Fertilized females disperse in search of host plants. Dispersal is mainly passive but females, as well as males, are able to fly against weak wind currents and respond to ol- ') Gall midges in the strict sense are gall inducers. _ Among Cecidomyiidae, gall midges sensu lato, some aberrant forms are predators. These are not sedentary. factory cues (McKay & Hatchett, 1984; Skuhra- vy et al, 1983; Sylvén, 1970). Eggs are usually laid on or close to the site where the neonate larva will penetrate the host plant. There is a considerable variation in clutch-size. The number of eggs may be one per Oviposition or up to five. Some species, e.g., Contarinia pulchripes (Kieffer), deposit all (up to 150) eggs in one batch (Parnell, 1963). Lar- vae, eclosed from the same clutch, are gregari- ous within a gall. Many midge species produce unisexual fami- lies, 1.e., the offspring of one female are either all male or all female. This mechanism of sex regulation might be common in gall midges be- cause the sex ratio departs in many cases from 1:1, the ratio expected in obligatory crossbreed- ing species. The mechanism has been studied by Metcalfe (1935) and Gallun & Hatchett (1969) for the Hessian fly, Mayetiola destructor (Say). Characteristics of the host plant, such as chemical composition and phenology, may have an important impact on gall midge development and, ultimately, on fitness. Host plants that are selected for oviposition may be less suitable, or even unsuitable for larval development. Females of Dasineura brassicae (Winnertz), for example, prefer pods of Brassica napus and B. campestris for oviposition but also lay eggs on B. juncea and B. nigra. However, the percentage of hatched eggs on the latter pair of host species is lower and larval development less successful, resulting in females with lower egg production (Ahman, 1981 and in press). Females of Haplo- diplosis marginata (Von Roser) search first for grasses or cereals, but if these are not available, especially during outbreaks, they will lay eggs upon any other plant and even on the soil. However, galls are only induced in grasses be- longing to the tribes Triticeae and some Ave- neae. Many eggs are laid upon Avena sativa, but there is very little survival on this species. For that reason Avena sativa is suggested for bio- logical control of Haplodiplosis in schemes of crop rotation (Skuhravy et al., 1983). Another factor for successful larval devel- opment is synchronization of host plant and gall midge phenologies. Winter varieties of wheat and barley are less susceptible for Haplodiplosis than summer varieties because neonate larvae are unable to penetrate, at the time of attack, the more mature tissues of earlier planted varieties (Nijveldt & Hulshoff, 1968; Skuhravy, 1982; Skuhravy et al., 1983). Phenological synchroni- zation is also important in other gall midge 196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 pests, e.g., Orseolia oryzae (Wood-Mason) on rice (Van Vreden & Arifin, 1977), Thecodiplosis brachyntera (Schwägrichen) on pine trees (Skuhravy & Hochmut, 1969; Skuhravy, 1970). Hatchett & Gallun (1970) demonstrated a ge- netic basis for the ability of Mayetiola destruc- tor (Say) to survive on different races of wheat. These races of wheat possess genes for resis- tance against attacks of Mayetiola, which on its turn can also be subdivided into races having genes to overcome this resistance. The gene-for- gene interaction between wheat and Hessian fly might have been developed as a reciprocal pro- cess (Gallun, 1977). At the end of this section on life history pat- terns and host plant suitability we may conclude that variation in life history patterns, relevant for particular modes of speciation, is mainly present during the free-living adult phase. Sometimes a highly coupled niche structure ex- ists indeed: if the pupation site is the gall, and the host plant is long-lived, emerged females may mate and lay eggs on the same host plant. Female dispersal is especially reduced when the eggs are laid in few (or only one) large batches (Weis et al., 1983). However, swarming of molles and daeaimesion of as by the mecha- nism of unisexual families considerably reduces the possibilities of assortative mating and hence sympatric speciation. Furthermore, oviposition on alien host plants occurs under some condi- tions, but the possibility of larval development may be a considerable hurdle for host plant shifts. Therefore, allopatric models of speciation will prevail in gall midges. Nevertheless, it is hard to imagine that in genera which exhibit ex- treme resource partitioning, such as the 62 Ste- faniola species on Haloxylon, or the 28 Rhopa- lomyia species, distinguished by Jones et al. (1983) on Artemisia tridentata, have exclusively radiated according to an allopatric model of speciation. TAXONOMY OF GALL MIDGES Gall midges belong to the nematoceran fami- ly Cecidomyiidae (4,300 described species according to estimates of Skuhravä, in press). Within the suborder Nematocera the Cecido- mylidae are a distinct group: wing veins are generally weak and reduced in number, the cos- tal vein is usually continuous around the wing and tibial spurs are absent. The larvae, usually bright yellow, orange or reddish in colour, pos- sess a supernumerary “neck” segment between head and thorax, which allows a great mobility of the head. On the ventral side of the protho- rax a peculiar sclerotized plate, the sternal spat- ula, is usually present. According to Mamaev (1968) the Cecidomyiidae are closely related to the mycetophagous scavengers Sciaridae, Sca- topsidae and Hyperoscelidae. A generally adopted subdivision of the family is still lacking. Mamaev (1968), following Rub- saamen & Hedicke (1925—1939) distinguished only two subfamilies: the Lestremiinae, with undifferentiated tarsi and with the ocelli usually present; the Cecidomyiinae with short first tar- someres and ocelli absent. Möhn (1955), fol- lowed by many modern students of the group, proposed a third subfamily Porricondylinae. However, he was only able to separate this sub- family by larval characters concerning position and shape of the anal aperture. Mamaev, refin- ing Rübsaamen & Hedicke’s system, differ- entiated the system to the subtribal rank, but many other specialists consider his system ten- tative and do not use it. The system used by Skuhrava (in press) in her catalogue is compared here with Mamaev’s system in table 1. Further differences concerning the Cecidomyiinae deal with taxa that are difficult to place. First, the Stomatosematidi in Skuhrava’s system, for ex- ample, share various archaic traits, such as wing venation (Rs well developed) and female genita- lia (short, not extensile, sometimes even two- segmented cerci) with Porricondylinae, but dif- fer from this subfamily by the male genitalia, which are reminiscent of those of Lasiopteridi. Gagné (1975), therefore, proposed an indepen- dent supertribal status for this taxon. Secondly, Gagné (1976) placed Oligotrophini and Lasiop- terini in the supertribus Lasiopteridi because these tribes share derived character states of fe- male genitalia and antennal flagellomeres; these are lacking in their sister-group Ledomyiini, which in its turn is characterized by derived conditions regarding tarsal claws and male geni- talia. The relationships of Brachineura, Epimyia and Rhizomyia, placed in separate tribes, are still unclear. These genera are now placed in La- siopteridi, but may be better regarded as un- placed (Gagné, 1976). Because of the still very uncertain relationships of Gagné’s Stomatose- matidi and Ledomyiini, we here adopt Ma- maev’s classification, at least as far as it concerns the tribal subdivisions. Our special attention is focussed on host plant relations and their importance for the evo- lution of the gall midges. Therefore we will now analyze the phyletic relations of the tribes in Roskam: Gall midge — host plant associations 197 Table 1. Comparison of systems of Cecidomyiidae according to Mamaev (1968) and Skuhravä (in press, pre- sented with permission from the author). I, inquiline; M, mycetophagous; P, phytophagous and gall inducing; Z, zoophagous. MAMAEV LESTREMIINAE 3 tribes, 7 subtribes CECIDOMYIINAE Heteropezini Porricondylini Lasiopterini 6 subtribes Oligotrophini Oligotrophina + 4 more str. Brachyneurina Rhizomyiina Epimyiina Stomatosematina Asphondyliini 3 subtribes Cecidomyiini SKUHRAVA LESTREMIINAE 2 supertribes, 8 tribes PORRICONDYLINAE Heteropezini + Leptosynini Porricondylini + 7 more tribes CECIDOMYIINAE Lasiopteridi Lasiopterini no subdivision Oligotrophini Ledomyiini Brachyneurini Rhizomyiini Epimyiini Stomatosematidi Asphondyliidi 4 tribes Cecidomyiidi no subdivision species feeding mode 13 subtribes connection with their feeding modes. All Ceci- domyiinae share the absence of ocelli and the shortening of the first tarsal segment of legs, both derived character states. The feeding mod- es (table 1) in this subfamily are most diverse, ranging from mycetophagy to various forms of phytophagy and zoophagy (Mamaev, 1968). All gall inducing midges, the “true” gall midges, be- long to the Cecidomyiinae. Heteropezini and Porricondylini, with primitive wing venation (Rs usually present) and larval morphology (pattern of setae on the final two abdominal seg- ments and location of the anal aperture), are mycetophagous, as are all Lestremiinae and all forms of the related families Sciaridae, Scatopsi- dae and Hyperoscelidae. Therefore, feeding on decaying organic material must be regarded as the original feeding mode of Cecidomyiidae (Southwood, 1972; Mamaev, 1968; Roskam, in press). Mycetophagy is also the feeding mode of oligotrophine Rhizomyiina and some species of Ledomyia. Although the larvae of Brachineuri- na, Epimyiina and Stomatosematina are un- known, these are expected to be mycetophagous too (Mamaev, 1968; Gagné, 1975). Furthermore larvae of the oligotrophine genus /sogynandro- myia live in the upper layer of forest soil (Spungis, 1981). Mycetophagy i is also common in the tribe Ce- cidomyiini; it is the feeding mode of Buhro- myiella, Camptodiplosis, Clinodiplosis, Dichae- tia, Dichodiplosis, Echinella, Giardomyia, Karshomyia, Mycetodiplosis, Mycocecis, Myco- diplosis, Neoisodiplosis and Neomycodiplosis, 59 species together. Some of these genera are close- ly related, e.g., Möhn’s (1955) “Mycodiplosis group” and “Clinodiplosis group”. Mamaev (1968) considered, on morphological criteria, mycetophagous Oligotrophini and Cecidomyii- ni primitive forms within these two tribes. No 198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 — te = ta w = = = = 9 ui > > 3 = ® Ernie E pei Cc £ = en 9 ©) = O (©) 4 ic q D 5 © a D == re] (cb) (ep) O a] O < u) = 5 = a ty N > = 5 N T a > 2 Se © E 3 ©) des (©) © + je) = © = O D = ped do D) (4) O © _ © T Q O O 7 8 10 11 Fig. 1. Phylogenetic relations of the Cecidomyiinae tribes. Black squares indicate synapomorphic conditions. 1, ocelli absent, shortened first tarsomeres; 2, paedogenesis; 3, larval anus shifted into ventral position and slit-like; 4, reduction in number of Malpighian tubes of larval digestive tract, reduced number of dorsal and ventral papil- lae on the larval eighth abdominal segment; 5, substitution of parameres in male genitalia by mediobasal out- growths of basimeres sheathing aedeagus, retractile ovipositor with fused cerci; 6, mediobasal outgrowths of male genitalia absent, number of adult antennal segments basically fixed, 2 + 12; 7, constriction in larval mid- gut shifted proximally; 8, wing vein R, closely adjacent to R, and C, reduced in length, antennal segments not or barely sexually dimorphic and barrel-shaped, characteristic ovipositor with hooks and spines adapted to abrade plant tissue; 9, binodal male antennal segments with looped circumfila, reduction of the eighth tergite of the female abdomen; 10, loss of the constriction in the larval mid-gut; 11, necks of antennal segments reduced in length with reticulate, closely appressed circumfila, retractile needle-like ovipositor with fused (reduced?) cerci. For further explanation see text. mycetophagous representatives are known of mites, six predate on aphids, five on coccids and Asphondyliini and Lasiopterini. two attack other cecidomyiids. Three genera are Zoophagy is mainly restricted to Cecidomyii- endoparasitoids of aphids and psyllids. Some of ni. Four genera are known as predators of these zoophages are important agents in biolog- Roskam: Gall midge — host plant associations 199 Table 2. Tribal preference for vegetative (veg.) and generative (gen.) host plant tissues. Gall midges belonging to the “mixed” category attack both types of tissues. Data are from an analysis of keys on plant galls by Buhr (1964—1965), only described gall midge species included and inquilines excluded. Expected values according to “chi-square” calculation (in brackets). TRIBUS 1. LASIOPTERINI Lasioptera Ozirhincus Stefaniella 3 monotypic genera TOTAL TRIBUS 2. OLIGOTROPHINI Arnoldiola Bayeria Cystiphora Dasineura Geocrypta Iteomyia Jaapiella Janetia Janetiella Lathyromyza Macrolabis Mayetiola Misospatha Neomikiella Oligotrophus Physemocecis Rabdophaga Rhopalomyia Wachtliella 21 monotypic genera w w 0 0 8 0 0 1 2 0 1 0 0 0 (0) 4 al w TOTAL 216(186.3) 54 (85.7) TRIBUS 3. ASPHONDYLIINI Asphondylia Placochela Polystepha 2 monotypic genera TOTAL 23 (10,0) TRIBUS 4. CECIDOMYIINI N w Ametrodiplosis Antichiridium Contarinia Diodaulus Harmandia Loewiola Macrodiplosis Massalongia Planetella Plemeliella Thurauia Tricholaba 18 monotypic genera 0 N & D ND © N ND D À N O1 Na N (o>) a D r en WONNWNNN & © Ho O05o05oION H ive} r oa N TOTAL 75 (91.5) 60 (42.1) © ND NI hH © © © À © © © H © w © H WOO © 200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 ical control. Apart from the Cecidomyiini, some species of Trotteria (T. galıı Rubsaamen and T. ligustri Barnes), Lasiopterini, and of Ledomyia (L. acarıphaga Marikovskij, L. acerina Giraud and L. cardui Kieffer), Oligotrophini, are sup- posed to be predators. Other forms of both gen- era are inquilines, and are regarded as early off- shoots within their respective tribes (Mamaev, 1968; Mohn, 1966). No conclusion is possible about the origin and evolution of the zoopha- gous Cecidomyiini. They may have evolved polyphyletically from either mycetophagous or phytophagous ancestors. Mohn (1955) indicated three groups of related genera, namely, the Les- todiplosis group (e.g., Lestodiplosis, Feltiella and Therodiplosis, predators of gall midges, aphids and mites), the Phaenobremia group (e.g., Phaenobremia, Aphidoletes and Monobremia, on aphids) and the mutually close endoparasi- toids Endaphis and Endopsylla. A cladogram of Mamaev’s tribal division of the Cecidomyiinae is presented in fig. 1. The synapomorphies (shared derived character states) 5 and 6, concerning male genitalia and antennae of both sexes, validate the two com- plexes of Oligotrophini-Lasiopterini and Ceci- domyiini-Asphondyliini. At dichotomy A, a la- siopterine form adopted phytophagy and sepa- rated from the Oligotrophini. Then, at a further dichotomy an oligotrophine form, becoming phytophagous too, separated from the remain- ing mycetophagous Oligotrophini. A similar process, starting at B, subsequently gave rise to phytophagous Asphondyliini and Cecidomyii- ni. Hence, unless mycetophagy in Oligotrophi- ni and Cecidomyiini is a derived feeding mode, the transition to phytophagy, culminating in gall inducing, occurred several times in a paral- lel way. Morphological arguments (fig. 1, the synapomorphies 7, 8, 10 and 11) as well as argu- ments emerging from gall midge parasitoids make a secondary transition to mycetophagy highly improbable. Mycetophagous and phy- tophagous cecidomyiids both have scelionid parasitoids. Chalcidoid parasitoids, however, are abundant on phytophages but do not attack mycetophages. If mycetophages have evolved from phytophages, undoubtedly some chalcı- doid parasitoids would have followed their hosts and would now be present on myceto- phages (Mamaev, 1968). SPECIALIZATION ON HOST PLANT ORGANS The transition from mycetophagy to phyto- phagy is supposed to coincide with the expan- sion of the angiosperms during the Upper Cre- taceous, about 65 million years ago (Klausnitz- er, 1977; Mamaev, 1968; Zwolfer, 1978). In Upper Miocene formations, 30 million years ago, all gall midge tribes were well represented (Gagné, 1973). Two prerequisites presumably were responsible for angiosperm expansion, namely, the progressive development of the conducting system ensuring intensive movements of sap and rapid progress in the de- velopment of the flower as an adaptation to in- sect pollination (Takhtajan, 1954). If the earlier assertion of a polyphyletic transition to phyto- phagy is true, it might be reflected in different specializations of the phytophagous members of the various tribes to the different progressive developments of their host plants. In other words, some tribes might basically be adapted to exploit the conducting system of their host ~ plants, subsequently colonizing other organs, such as leaves, whereas other tribes might be primarily adapted to generative structures, such as flowers, compact inflorescences as heads of Asteraceae and fruits. With the help of table 2 we can investigate whether differences exist at the tribal rank in the specialization of gall midge species on tissues of their host plants. The species, compiled from Buhr (1964—1965), are subdivided into three categories: those causing deformations of vege- tative structures, of generative structures and those with a “mixed” strategy, attacking both vegetative and generative structures. The data have been submitted to a chi square-test. The null hypothesis, i.e., no significant differences exist between tribes regarding specialization for organs of host plants, has to be rejected (x2 = 54.8, DF = 6, P<<0.001). The frequencies of Lasiopterini accord with the expected ones (in brackets), although the species of this tribe have ovipositors which are primarily adapted to abrade stems and to insert eggs into them. Ozirhincus, although sharing these morphological features, is aberrant, gal- ling generative instead of vegetative tissues. In North America, where the Lasiopterini are well represented, most species are stem feeders since only six out of 70 species belong to the “genera- tive” or “mixed” category (Felt, 1940; Gagné, 1969). Oligotrophini have a distinct preference for vegetative organs. Aberrant oligothrophine genera are Gephyraulus, Kaltenbachiola and Se- mudobia; aberrant species are found in most larger genera, viz., Dasineura, Jaapiella, Macro- labis, Misospatha, Rhopalomyia and Wachtliel- Roskam: Gall midge — host plant associations 201 Table 3. Distribution of gall midge species among orders of vascular plants. Only orders with West Palaearctic representatives have been considered and are subdivided into a fundamentally woody (+) category and a funda- mentally herbaceous one (Hutchinson, 1969). The numbers of the orders refer to Takhtajan (1980), the numbers of plant species are taken from Rothmaler (1972), those of gall midge species from Buhr (1964—1965). (1) = Hutchinson’s Brassicales; (2) = Hutchinson’s Umbellales. (sub)class plant midge species species Lycopsida Lycopodiales Selaginellales Isoetales Sphenopsida Equisetales Pteropsida Ophioglossales Osmundales Polypodiales Marsileales Salviniales Ginkgoopsida Ginkgoales Taxopsida Taxales Coniferopsida Pinales Magnoliidae Aristolochiales Nymphaeales Ranunculidae Ranunculales Papaverales Hamamelidae Hamamelidales Urticales Fagales Myricales Juglandales Caryophyllidae Caryophyllales Polygonales Plumbaginales Dilleniidae Paeoniales Theales Violales Capparales 1) Tamaricales Salicales Ericales Primulales Malvales + Euphorbiales + Thymelaeales r > Saxifragales Rosales Fabales Myrtales Rutales Sapindales Geraniales Polygalales Cornales Araliales 2) Celastrales Santalales Rhamnales Qn 3 7 7 6 5 1 5 7 2 5 7 9 3 1 7 3 1 1 1 to Asteridae Gentianales Oleales Dipsacales Polemoniales Lamiales Scrophulariales Campanulales Asterales Alismidae Alismatales Najadales Liliidae Liliales Orchidales Juncales Cyperales Poales Arecidae Typhales Arales TOTALS woody orders herbaceous orders 202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 la. Asphondyliini occur predominantly on gen- erative structures. However, Polystepha is aber- rant as are nine out of 28 Asphondylia species. The situation is rather complex in Cecidomyii- ni. Although almost half of the species induce galls in vegetative tissues, a substantially larger portion is predicted. Contarinia, by far the largest genus of Cecidomyiini, is almost com- pletely responsible for deviations of this tribe from the expected value. The ambiguous preference of Cecidomyiini does not allow drawing conclusions about the original specialization of the group. Two alter- native ways of radiation might be possible. First, as in Oligotrophini, a phytophagous be- haviour started on vegetative parts and differ- entiation at the generic level coincided with a shift to generative parts. The radiation of Con- tarinia has than to be regarded in connection with this shift from vegetative to generative or- gans. Secondly, as in Asphondyliini, the prefer- ence of Contarinia for generative organs should be regarded as original. Specialists of vegetative tissues, belonging to Contarinia as well as to other genera, then have ancestors on generative parts. Synapomorphic conditions of male geni- talia and antennae in Cecidomyiini and Asphon- dyliini (fig. 1: 6) support the second alternative, but a further phylogenetic study (according to current opinion Contarinia is not monophylet- ic!) is needed to solve this problem. SPECIES RICHNESS AND HOST PLANT DIVERSITY A further consequence of the assumed coin- cident radiation of gall midges and host plants is that these ecologically linked groups of organ- isms are expected to illustrate Eichler’s rule (Eichler, 1948; Price, 1977): host plant taxa with many species will support more midge spe- cies than taxa which are less diverse, because there is more scope for radiation among the - midges. In table 3 the accumulations of gall midge species are given for all orders of West Palaearctic vascular plants. All dicotyledonous 2.0 T = trees ,n=21 H = herbs, n = 35 fami © © 8 1.5 (7) te) 2 = Y 2 © a 1.0 (7) ® = T 5 £ T T m 0.5 H T H He mmh nd le || mm huhu _ (o) 0.5 1.0 1.5 2.0 2.5 plant species (log scale) Fig. 2. Interdependence of numbers of plant and gall midge species per plant family. Midge species according to Buhr (1964— 1965), plant species according to Rothmaler (1972). Roskam: Gall midge — host plant associations 203 orders which include ten species or more have host plants attacked by gall midges. Fagales, Salicales and Fabales (Leguminosae) in particu- lar have host plants which shelter numerous gall midge species. Large monocotyledonous orders, such as Juncales and Orchidales, as well as most ferns and gymnosperms, lack gall midges. Poales and Cyperales, however, have many rep- resentatives with gall midges. In fig. 2 the interdependence between the number of gall midge and host plant species per plant family is analyzed. Contrary to earlier stu- dents of insect species richness (e.g., Lawton & Price, 1979; Fowler & Lawton, 1982), we used the plant family and not the plant genus as the variate for “taxonomic diversity” (= taxonomic isolation) of host plants. This is because the pre- sent study involves many host plant families, whereas Lawton, Price and Fowler only dealt with one family, Umbelliferae. Furthermore, in different families different criteria are used to delimit genera, which makes genera unsuitable for comparison when more families are in- volved. For logarithmic transformed data taxo- nomical interdependence alone explains 52.5% (r = 0.72) of the variation in gall midge species richness on host plants, and is therefore a very important factor. When host plants are subdi- vided into woody and herbaceous categories the percentages are even higher, namely, 66.1% (r = 0.81) for woody and 63.3% (r = 0.80) for herbaceous host plants. These high percentages mean that taxonomic diversity, reflecting diver- sity of host plant chemistry (Hegnauer, 1962— 1973) is not the only but apparently a major fac- tor determining accumulation of gall midge spe- cies on host plants. Similar suggestions were made by Claridge & Wilson (1981), dealing with mesophyll-feeding leafhoppers. Price (1977) observed a similar interdependence for another group of endophytophages: Agromyzi- dae. In his calculation 50.4% of the variation of leafminer species per host plant family was ex- plained by the number of plant species in that family (r = 0.71). The regression lines for woody and herba- ceous host plants do not differ significantly by slope, only by intercept. Hence, gall midge spe- cies are more numerous in plant families with woody representatives than in those with herba- ceous ones: the “high apparency” of long-lived woody host plants (Fox, 1981; Lawton, 1983; Lawton & Schréder, 1977; Klausnitzer, 1977) also works positively out for gall midges. When the gall midges are subdivided into Table 4. Tribal preference with respect to the life form of host plants. Data after an analysis of the keys by Buhr (1964—1965). percentage of life form species number annual/ biennial perennial shrubs/ Lasiopterini Oligotrophini Asphondyliini Cecidomyiini tribes and host plants categories according to their life form (table 4), most gall midges of all tribes occur on perennial herbs, whereas short- lived herbs are poorly represented. The high score of Lasiopterini for short-lived herbs is mainly caused by one species, Lasioptera caro- phila F. Loew, which attacks many short-lived umbellifers. When analyzing the life form pref- erence of Lasiopterini for North America, where L. carophila is absent (compilation of Felt, 1940, and Gagné, 1969), 7.1% of 70 spe- cies occur on annual and biennial host plants, 67.1% on perennial herbs and 25.7% on shrubs and trees, values conforming to those of tribes other than Lasiopterini in Europe. The short-lived host plants need a further analysis. Many of these plants are characterized by conspicuous chemicals as furanocoumarins (Apiaceae) or mustard oil glucosides (Brassica- ceae). Short-lived Apiaceae are hosts for two polyphagous species, viz., Lasioptera carophila F. Loew and Kiefferia pimpinellae (F. Loew). Short-lived Brassicaceae harbour polyphagous Contarinia nasturtu (Kieffer). Dasineura brassi- cae (Winnertz) and D. sisymbrii (Schrank) and Gephyraulus raphanistri (Kieffer). Mayetiola destructor (Say), Haplodiplosis marginata (Von Roser) and Hybolasioptera cerealis (Lindeman) have many annual cereals in their host ranges. These cereals occur in high densities, in “flocks”, and germinate not far from the place where the previous generation lived. In this way they are “predictable” resources and resemble perennials. Finally, short-lived host plants are present among Chenopodiaceae (Haloxylon), Asteraceae (e.g., Senecio, Sonchus, Cirsium and Carduus) and Leguminosae (e.g., Lathyrus, 204 Lens, Medicago, Melilotus, Pisum and Vicia). These host plants either occur under natural conditions in dense populations, or are also cul- tivated. PARALLEL PATTERNS IN GALL MIDGE AND HOST PLANT EVOLUTION If related parasites live on related hosts, allo- patric speciation patterns in both groups of organisms may have evolved along parallel lines: dichotomies in host cladograms then have corresponding dichotomies in cladograms of parasites. Corresponding dichotomies or co- cladogeneses may be the result of a reciprocal process between hosts and parasites: parasite at- tack, reducing fitness of the host, provokes the host to develop defense or avoiding mecha- nisms. Parasites, on their turn, try to overcome host defenses by counter adaptations and so on. However, long term reciprocal interactions (de- fined by Janzen (1980) as co-evolution) are not the only process resulting in parallel patterns. Moreover, when they do so, they may be diffi- cult to measure. Parasites usually share their host plants with many other parasites, each pos- sessing different trophic links with their hosts (Klausnitzer, 1977). A change of a host, to avoid one parasite, might be advantageous for another. The complexity of interactions reduces the profits of that change (Fox, 1981). Changes in the host plant may also, and more frequently, be the result of responses to abiotic changes of the host plant habitat. Parasites may follow the changes of their hosts for their own benefit. This type of parallel evolution has been defined by Jermy (1976) as sequential evolution. Fi- nally, speciation processes in host plants and parasites may coincide, but as independent re- sponses to the same abiotic factor. Vicariance, caused by the same geographic isolation in sub- groups of hosts and parasites, may so cause a parallel pattern in the phylogenies of both groups (e.g., Roskam, 1979). Parallel patterns need not necessarily be strict because phytophages, unlike many parasites of vertebrates, have a free phase during their life- cycle. While dispersion of vertebrate parasites usually occurs by conspecific contacts of their hosts, dispersion of phytophages, at least in gall midges is possible during a free-living phase, as was reported in the above. They may shift to other, usually related, host species during that phase, causing disturbances of parallel patterns (Regenfuss, 1978). Whereas the host range of zoophagous and TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 saprophagous cecidomyiids is relatively wide (Nijveldt, 1969; Skuhrava, 1973), most gall in- ducing and inquiline species have only narrow host plant ranges. They usually attack some re- lated species belonging to the same genus, or species belonging to closely related genera. Ex- ceptions are, e.g., Lasioptera carophila F. Loew and Kiefferia pimpinellae (F. Loew) on Apia- ceae; Dasineura sisymbri (Schrank), D. brassi- cae (Winnertz) and Gephyraulus raphanistri (Kieffer) on Brassicaceae. Both host plant fami- lies are distinct by chemical compounds, fura- nocoumarins and mustard oil glucosides, re- spectively. Some Asphondylia species alternate host plants during their life-cycle, as do aphids. According to Orphanides (1975), the winter generation of the carob gall midge, Asphondylia sp., induces galls in pods of carobs, Ceratonia siliqua. Summer generations, however, attact : various, not related, plant species, viz., Capsı- cum and Solanum (Solanaceae), Capparıs (Cap- paridaceae), Eruca and Sinapis (Brassicaceae), Hypericum (Hypericaceae), Verbascum (Scro- phulariaceae), Sesamum (Pedaliaceae) and even monocots, viz., Urginea and Asphodelus (Lilia- ceae). It is still uncertain whether midges reared from these plants will be conspecific. Some, however, certainly are. A similar situation seems to exist in the soybean gall midge, As- phondylia sp., overwintering in soybean pods but with unknown summer hosts (Yukawa et al., 1983). Among gall midge genera large differences exist regarding the breadth of their host plant spectrum. In table 5 gall inducing midge genera are subdivided into three categories, namely, monophagous, oligophagous and polyphagous genera. Genera with eight species or more are listed, whereas smaller genera only are indicated by their number of species. The large genera Dasineura, Contarinia, Jaapiella and Macrola- bis, but also the smaller Wachtliella, are pre- sented in brackets, because they are highly artı- ficial and therefore do not allow conclusions about the affinities of their host plants. Interde- pendence between gall midge species diversity and breadth of the host plant spectrum seems to be absent: not only large genera as Asphondylia, Rhopalomyia and Lasioptera are polyphagous, but also many small genera consist of species which occur on host plants belonging to differ- ent families. Monotypic polyphagous genera are absent. Lasioptera and Neolasioptera are two large genera which are thought to be natural. Gagné Roskam: Gall midge — host plant associations 205 Table 5. Host plant spectrum of gall midge genera. M, monophagous genera, all host plants belong to only one genus; O, oligophagous genera, host plants belong to one family; P, polyphagous genera, host plants belonging to several families. Data from Skuhrava (in press, with permission from the author). For further explanation, see text. Dasineura Contarinia 151 Stefaniola 69 Asphondylia 54 Rhopalomyia 49 Lasioptera 45 Rabdophaga 38 Halodiplosis 37 Jaapiella 31 Macrolabis 31 Planetella 26 Mayetiola 25 Baldratia 23 Janetiella 15 Ametrodiplosis 13 Cligotrophus 10 Arnoldiola 8 Wachtliella 8 2 genera 7 4 genera 6 5 genera 5 3 genera 4 15 genera 3 18 genera 2 total natural genera (1969) revised the Nearctic species, of which Felt (1940) presented the host plants. Both gen- era are well represented on host plants belong- ing to the subclasses Rosidae (orders: Rosales, Fabales, Cornales and Rhamnales) and Asteri- dae (orders: Lamiales, Scrophulariales and As- terales). They are absent from Monocotyledo- nae; two species of Lasioptera occur on Ephe- dra (Gymnospermae, Gnetales). Lasioptera has five species on host plants of the subclass Ham- amelidae (Humulus and Quercus), from which subclass Neolasioptera is absent. On the other hand, Neolasioptera is represented in the sub- classes Magnoliidae (Lauraceae: Benzoin) and Ranunculidae (Ranunculaceae: Clematis) where Lasioptera is absent. Although both genera have accumulations of species on Rosidae and Asteri- dae, apparent parallel patterns with the phylo- genus nr. of species with more than one species geny of host plants belonging to these sub- classes are still lacking. Some smaller genera also have species attack- ing hosts belonging to unrelated families or even have species with a non-cecidogenic feeding mode. Janetiella, for example, occurs on hosts belonging to Pinaceae, Cupressaceae, Fagaceae, Ulmaceae, Chenopodiaceae, Brassicaceae, Le- guminosae, Vitaceae, Euphorbiaceae, Labiatae and Asteraceae. Host plants of Ametrodiplosis belong to ten families; two species are inqui- lines. Even among genera with only two includ- ed species, nine occur on host plants which are taxonomically distant. Physemocecis hartıgı (Liebel) causes galls on Tilia (Tiliaceae), where- as P. ulmi (Kieffer) occurs on Ulmus (Ulma- ceae). Antichiridium caricis Kieffer and A. stria- tum (Rübsaamen) cause galls on Carex (Cype- 206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 raceae) and Molinia (Poaceae), respectively. Plemiella abietina Seitner and P. betulicola (Kieffer) have Abies (Pinaceae) and Betula (Be- tulaceae), respectively, as host plants. These genera may involve examples of ecological op- portunists because the host plants on which their species occur share the same habitat. Other genera with two species, however, occur on host plants which have neither taxonomical, nor apparent ecological connections, e.g., Dic- tyomyia navasiana Tavares and D. salsolae Ta- vares on Santolina (Asteraceae) and Salsola (Chenopodiaceae), Schizomyia galiorum Kieffer and S. tami Kieffer on Galium (Rubiaceae) and Tamus (Dioscoreaceae). On the other hand, some larger genera ra- diated on closely related host plants. Stefaniola and Planetella have host plants belonging to on- ly one genus, Haloxylon and Carex, respective- ly. Rabdophaga occurs on Salicaceae (mainly on Salix) and Mayetiola on grasses. Baldratia and Halodiplosis exclusively occur on Chenopodia- ceae. When apparently monophyletic genera are taken together no less than 63% (38 out of 60 genera) radiated on host plants which are taxo- nomically close. Examples of parallel cladogenesis in gall midges and host plants will primarily be found in genera that radiated on taxonomically related host plants. In the next paragraph of this section some of these examples will be treated. The ex- amples are arranged according to the taxonom- ical rank of the host plants at which the radia- tion occurred. A. Host plant family Host plants of Asphondylia mainly belong to two families which are phylogenetically distant, namely, Leguminosae and Labiatae. Correlated with the taxonomic position of the host plants there is a specialization with respect to the host plant tissue. Out of 28 species mentioned in Buhr (1964—1965), all species on Labiatae (6) make flower galls, whereas 18 species on Legu- minosae are specialized on pods (11) or vegeta- tive parts (6); A. sarothamni H. Loew on Saro- thamnus causes galls in pods, flowers and shoots. Hence, species causing flower galls on Labiatae and fruit galis (and later in evolution- ary time, shoot galls?) on Leguminosae may represent two different evolutionary lines in this polyphagous genus. Three genera, Mayetiola (Oligotrophini), Haplodiplosis (Cecidomyiini) and Hybolasiop- tera (Lasiopterini), of which the latter two are monotypic, induce galls in culms and shoots of Poaceae. As a rule, they attack many wild grass- es. Some species, however, are extremely im- portant pests of cereals. Mayetiola destructor (Say), the Hessian fly, is the most important gall midge species damaging cereals. Wild grasses from which the species has been reported be- long to Cynodon (Poaceae-Eragrostideae), Phleum, Aegilops, and Agropyron (Poideae). Other Mayetiola species, usually one per plant genus, induce galls in culms and shoots of Ave- na, Brachypodium, Calamagrostis (various Mayetiola species occur in this genus), Dacty- lus, Holcus, Molinia, Phalaris, Poa and Secale (all Pooideae).- Giraudiella, one species, closely related to Mayetiola, induces galls on Phrag- mites (Pooideae). Hence, most host plants be- long to the subfamily Pooideae, but at a lower level apparent patterns are absent. B. Host plant tribe Four clusters of oligophagous genera, re- stricted to Asteraceae, are of particular interest with respect to parallel cladogenesis (table 6). All five species of Ozirhincus (Lasiopterini) in- duce fruit galls in host plants belonging to As- teroideae-Anthemidae, as does Lasioptera (Pro- lasioptera) niveocincta (Kieffer). The Nearctic genus Asteromyia (Lasiopterini), with 20 spe- cies, only induces galls in members of the tribe Asteraceae. Two related Oligotrophini genera, namely, Rhopalomyia (49 spp.), of which 14 are mentioned in Buhr, and Misospatha (5 spp.), are present in Anthemidae too. Cystiphora (6 spp.), which also belongs to the Oligotrophini, only causes galls in members of the subfamily Cicho- rioideae. In Cystiphora, there is host specificity below the genus level: C. hreracu (F. Low) and C. pilosellae Kieffer are restricted to the Archie- racium and Pilosella groups of species, respecti- vely. In Asteroideae-Cardueae both species of Loewiola (Cecidomyiini) induce leaf galls in Centaurea and Serratula, whereas Acodiplosis (1 sp.), close to Loewiola, is present on Inula (Inu- | leae). We may conclude that, contrary to gall midges occurring on grasses, midge genera on Asteraceae exhibit specificity at the tribal rank. C. Host plant genus Many gall midge genera are restricted to only one host plant genus. Sometimes, related midge genera have related host plants. Dryomyia, for example, with four species, is reported from leaves of Quercus, whereas its relative, Harti- giola, with one species, causes galls in leaves of Roskam: Gall midge — host plant associations 207 Table 6. Gall midge genera associated with Asteraceae. Subdivision of Asteraceae according to Engler (1964). | Cecido- Oligo- myiini trophini (Prolasioptera) Loewiola Acodiplosis Rhopalomyia Misospatha Cystiphora Ozirhincus Asteromyia subfamily L. Asteroideae Eupatorieae - Senecioneae Calenduleae = Anthemideae Anthemis Achillea Matricaria Chrysanthemum Tanacetum Artemisia Astereae Erigeron Aster Solidago Bigelowia Inuleae Inula Cynareae Serratula Centaurea Cichorioideae Cichorieae Hypochoeris Leontodon Scorzonera Chondrilla Taraxacum Sonchus Crepis Hieracium 208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 Fagus. Semudobia, with three Palaearctic spe- cies, occurs on Betula, whereas its relative Mik- omyia, with one species, causes galls in Corylus. Within this category three examples have been selected, viz., Rabdophaga (Oligotrophini), with 38 species on Salix, Planetella (Cecido- myiini), with 26 species on Carex, and Semudo- bia (Oligotrophini) in fruit catkins of Betula. One species of Rabdophaga is known from twigs of Populus, the other species cause galls in shoots, twigs and catkins of Salix. Within Salix, there seem to be three levels of specialization that coincide with the division of Salix into sub- genera. Infections are absent from the subgenus Chaematia Dumortier, all prostrate, small shrubs. Some Rabdophaga species occur in both remaining subgenera: Salix s.s. (trees and large shrubs) and Caprisalix Dumortier (shrubs). R terminalis Kieffer, for instance, occurs in shoots and leaves, R. rosaria (H. Loew) in shoots, R. deletrix (Rubsaamen) in buds and R. nervorum (Kietter) in leaves. However, a number of spe- cies exclusively attack willows of the subgenus Salix: R. saliciperda (Dufour) in twigs, R. trian- draperda Barnes in twigs, R. heterobia (H. Loew) in shoots and in male catkins. The major- ity of Rabdophaga species are restricted to the large genus Caprisalix. Table 7 presents the host plant relations of 13 Planetella species which are recorded in Buhr (1964—1965). Two groups of species are dis- tinct, namely, the species which cause galls in sedges belonging to both sections Vignea (Beauvois) Kükenthal and Carex, and those which are restricted to the section Carex. Re- cords are absent from the remaining subgenera Indocarex Baillon and Primocarex Kükenthal. When analyzing Rübsaamen & Hedicke’s (1925—1939) keys, there is a parallel situation in gall midges and host plants with respect to the state of derivativeness of some characters: the species which exhibit apomorphic character states in the shape of the adult thorax and/or number of male flagellomeres, viz., Planetella tarda (Rübsaamen), rosenhaueri (Rübsaamen), fischeri (Frauenfeld), tumorifica (Rübsaamen) and cornifex (Kieffer), only occur on sedges which in their turn share synapomorphies with respect to the differentiation of their inflores- cences in male and female spikes. Unfortu- nately, the two poorly known species P. kneuckeri (Kieffer) and P. subterranea (Kieffer & Trotter), which were only reported from sedges belonging to the section Vignea, are not mentioned in Rubsaamen & Hedicke. Table 7. Species of Planetella associated with subge- nera of Carex. Subdivision of Carex according to Chater (1980). caricis baudisi granifex arenaria subterranea gallarum frireni kneuckeri tarda cornifex rosenhaueri fischeri tumorificus Cladograms of gall midges and host plants have been provided by Roskam (1979) for Se- mudobia (five spp.), including two Nearctic species, and Betula (about 40 spp.) (fig. 3). Four dichotomies, or events of speciation, in Semu- dobia have corresponding branchings in Betula. First, S. skuhravae Roskam induces galls in the bracts of fruit catkins, whereas S. betulae (Win- nertz), S. tarda Roskam, S. brevipalpis Roskam and S. steenisi Roskam, sharing apomorphies of larval and adult morphology, make galls in fruits. This dichotomy 1 is reflected in Betula at the section level. Whereas birches belonging to the sections Costatae (Regel) and Humiles (Koch) have erect catkins with fruits overwin- tering in the trees, birches of the sections Excel- sae (Koch) and Acuminatae (Regel) bear pen- dent catkins and disperse their fruits in the au- tumn of the year of flowering, an apomorphic condition. Acuminate birches lack Semudobia galls. S. skuhravae causes galls in birches of all remaining sections, but the fruit galling midges are only present on birches of the section Excel- sae. The structure of the catkins in the latter Roskam: Gall midge — host plant associations 209 Zu sku COSTATAE\a2 o/ Lu LU D O HUMILES E (©) SEMUDOBIA BETULA bet ACUMINATAE A a 9 \& 2 A U > FAI tar populifoliasw coerulea-gr._2 platyphylla~w Ww z Al È Ww nee D | m 7 x papyrifera\w m) À 4 CU fontinalis > davuricasw — < a. pubescens,2 Fig. 3. Parallel cladogenesis in Betula and Semudobia. The numbers refer to corresponding dichotomies in the cladograms. The branch which is not supported by apomorphies is indicated by a question mark. CIRC, Cir- cumboreal; NE, Nearctic; PAL, Palaearctic; PUB, Pubescentes; VERR, Verrucosae; bet, Semudobia betulae; bre, S. brevipalpis; sku, S. skuhravae; ste, S. steenisi; tar, S. tarda. section allows fruit galling Semudobia species to hibernate in the soil, which is a favourable con- dition (Mohn, 1961). Dichotomy 2 in Semudobia is parallelled by Betula at the series level: S. tarda is common in birches of the series Pubescentes Sukaczew of Excelsae, whereas S. betulae predominates in birches belonging to the series Verrucosae Su- kaczew. This branching separates birches of dif- ferent habitat conditions and apparently evolved under allopatric conditions. However, the recent birches of both series may occur sympatrically, as do S. betulae and S. tarda. Both midge species are able to induce galls in birches belonging to both series, but their pref- erence is different, reminiscent to the original, allopatric situation (Roskam & Van Uffelen, 1981). Finally, there is a correspondence regarding the third and fourth branchings, as a result of geographical vicariance. In both series of the section Excelsae different species occur in the western and eastern part of both Palaearctic and Nearctic. In the “betulae group” of Semudobia species, viz., S. betulae, S. brevipalpis and S. steenisi, this vicariance is incompletely parallel- led: S. brevipalpis and S. steenisi being restricted to the East and West Nearctic, respectively, and S. betulae occurring in the whole Palaearctic (Roskam, 1979). We must conclude, as was expected in phyto- phages, that parallel branchings in Betula and Semudobia are not complete. Moreover, fruit- galling Semudobia species were able to shift to other phyletic lines of birches under circum- stances of secondary sympatry. Real reciprocal adaptations are absent. The first dichotomy 1s an example of sequential evolution: a change in the construction of the catkin, in favour of dis- persal of the birch fruits, is exploited by the fruit-galling midges to improve their conditions for hibernation. All other branchings evolved simultaneously in plants and midges under con- ditions of allopatry. 210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 3, 1985 CONCLUSIONS 1. Two feeding modes are common in Ceci- domyiidae, namely mycetophagy and phyto- phagy, the latter eventually culminating in gall induction. Phytophagy, occurring exclusively in Cecidomyiinae, must be considered as a derived feeding mode. Outgroup comparison of larval and adult morphology, and feeding modes of re- lated nematoceran families are arguments for this conclusion. Within the Cecidomyiinae two clusters of tribes can be defined on morphologi- cal synapomorphies, viz., Oligotrophini — La- siopterim and Cecidomyiini — Asphondyliini. Because in both clusters mycetophagous repre- sentatives exist, and a secondary transition from phytophagy to mycetophagy is highly improba- ble, an independent, and hence polyphyletic transition from mycetophagy to phytophagy must be accepted in gall midges. 2. When species richness of gall midges is de- fined for families of host plants (logarithmically transformed data), the taxonomic interdepen- dence of gall midge and host plant species num- bers explains more than half the variation of gall midge species richness on those plants. Long- lived, woody plants accumulate more gall midge species than short-lived herbs. Contrary to some results for external plant feeders, taxo- nomical diversity of host plants is an important variate for this group of endophytophages to explain their radiation. 3. At the tribal rank gall inducing Cecido- myiidae are differently adapted to structures of their host plants. In Lasiopterini and Oligotro- phini significantly more species are adapted to vegetative organs, such as stems, vegetative shoots and leaves, whereas in Asphondyliini and Cecidomyiini more species are adapted to generative organs, such as flowers, inflores- cences and fruits. If the preference for genera- tive tissues in Contarinia is original for Cecido- myiini, the different preference of gall midge tribes for host plant organs may function as evi- dence for a polyphyletic transition to phytopha- 4. Most gall inducing midge species have nar- row host plant spectra. Limits at the gall midge genus level are usually narrow too: species of the same genus have host plants which are also congeneric or belong to some closely related genera. Although examples of ecological oppor- tunists are in the minority, they also exist in gall midges. Gall midge — host plant relations may be diffuse: parallel traits between gall midge and host plant phylogenies are absent, due to shifts of gall midges to, usually, related host species during the free living adult phase. Sometimes re- markable parallel traits are present in gall midge and host plant phylogenies. Dichotomies at the species level in gall midges match dichotomies at various levels of host plant taxonomy. In As- phondylia a dichotomy is present at the host plant family level: one cluster of species causes galls in flowers of Labiatae, whereas another cluster is restricted to pods or vegetative parts of Leguminosae. Loewiola and Acodiplosis, two closely related Cecidomyiini, both occur on As- teraceae, but have host plants belonging to the different, also mutually close tribes Cynareae and Inuleae, respectively. In Rabdophaga and Planetella specificity is present below the genus level of host plants. Species of the latter genus, . which exhibit morphological synapomorphous character stages occur on sedges which in their turn are also characterized by synapomorphies, indicating parallel evolution of both groups. In Semudobia parallel traits with host plant phylo- geny are obvious. Some corresponding dichoto- mies evolved independently in both systems as a result of geographical isolation, one event of parallel cladogenesis apparently is the result of sequential evolution. 5. Prerequisites for sympatric speciation are present in gall midges which live in perennial plants, hibernating and pupating in the galls; mating and oviposition then occurs in close proximity to the gall. However, assortative mating, another prerequisite, is unlikely in many instances because of the production of unisexual families by females and swarming flights of virgin males. Furthermore, in cases of host shifts, even to closely related plant taxa, a considerable reduction of fitness can be ob- served. 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Evol.-forsch. 22: 211—233. bed al 6 da PA Fio ie 8 lo rt ae ni | Be S à KL E n ke | SUR om 4 x % peut) Waits À A x PRIE ue A parut re ni AQU ae RES OST I > te 5 À Ld ae FRA à At it lj DEEL 128 AFLEVERING 4 1985 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD R. H. COBBEN. — Additions to the Eurasian saldid fauna, with a description of fourteen new species (Heteroptera, Saldidae), pp. 215—270, figs. 1—21, maps 1—4. Tijdschrift voor Entomologie, deel 128, afl. 4 Gepubliceerd 20-XII-1985 hear gaar. he x EN è a Be Heidi Pr rende ft: ci APE, fe it ADDITIONS TO THE EURASIAN SALDID FAUNA, WITH A DESCRIPTION OF FOURTEEN NEW SPECIES (HETEROPTERA, SALDIDAE) by R. H. COBBEN Department of Entomology, Agricultural University, Wageningen, The Netherlands ABSTRACT The following new species of Eurasian Saldidae are described: Halosalda coracina (Greece), Saldula hasegawai (far East USSR, Japan), S. tarwanensis (Taiwan), S. sibiricola (USSR: Kazakhstan), Macrosaldula clavalis (USSR: Georgia), M. inornata (Iraq), M kerzhneri (USSR: Kazakhstan), M. koktshetavica (USSR: Kazakhstan), M. miyamotoi (Ja- pan), M. shikokuana (Japan), M. simulans (Siberia, Mongolia), M. violacea (Far East of USSR, Japan), Calacanthia grandis (China), Salda kiritshenkoi (USSR: Central Asia, Far East, N. E. China, Japan). A new subspecies M. oblonga acetabularis is described from Kazakhstan. Salda nevadensis Wagn. and S. littoralis piechockit Wagn. are synonymized with S. littoralis L. Lectotypes are designated for Salda micans Jak., S. splendens Jak. and Macrosaldula roborowski Jak. comb. n. (transferred from Chartoscirta). The Macrosaldula clade is discussed; it is provisionally taxonomically treated as a genus. A tentative key is presented for 21 species of Macrosaldula presently recognized in Eurasia. Preliminary rede- finitions of M. jakovleffi Reut. and M. nivalis Lindb. are provided, whereas the status of M. mongolica Kir. needs further confirmation. The known localities of Macrosaldula, Teloleu- ca and Salda species (except for those from W. Europe) are mapped and the zoogeography of these genera is briefly discussed. CONTENTS Miner ne TONNERRE ANT EEN 215 Description of species and comparative BOCES hrc ce AEN 217 Discussion of Eurasian Salda species ..... 246 Tentative key to Macrosaldula species .... 254 Comments on the zoogeography of Ma- crosaldula, Salda and Teloleuca........ 262 Ncknomledeementene ve a. 263 REE ante 263 INTRODUCTION The present study is based predominantly on a revision of saldid material collected by Rus- sian and Japanese heteropterists in the eastern parts of the Palaearctic. I have added some new species from Greece, Iraq and China, which certainly belong to supraspecific taxa having originated in the northern hemisphere of the Old World. The description of one species from Taiwan is included here as well, although it may be a member of a species group from a more southern origin. Although the description of some species is based on only scanty material and, consequently, the knowledge of variability 215 and distributional patterns is limited, I refrained from a further delay of publication. This revi- sion may stimulate the study of material I have not seen, and exploration of areas from which no or only sparse data are available. Detailed ecogeographical analyses of population struc- tures of selected species groups, as for example undertaken by Dr P. Lindskog (in prep.) on the complex S. orthochila-burmanica (see p. 225), are dependent on more numerous material than I had the opportunity to study. The present paper does not include a revision of the abundant material in Russian and Japa- nese collections of small-sized typical Saldula and Micracanthia species. Such a revision is ur- gently needed in order to understand the zoo- geography of these world-wide genera with preponderance of species occurring in the northern hemisphere. I sincerely hope that Dr N. N. Vinokurov (Yakutsk, USSR), who initi- ated a fine, detailed study on this difficult group of saldids in Eastern USSR (Vinokurov, 1975, 1978, 1979a—c, 1981), will eventually be suc- cessful in preparing a comprehensive revision. We may expect more examples of Holarctic dis- 216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Fig. 1. a—g, Saldula hasegawai. a, paramere; b, grasping plate of 3; c, apex of paramere; d, base of penisfilum of two specimens; e, parandria; f, subgenital plate; g, median ardosomal sdeste. h—p, Halosalda. h, o, p, H. lateralis; i, —|, H. coracina; m, n, H. concolor; h, i, paramere, left figure of i as seen in direction of arrow in right figure; j, penisfilum; k, grasping plate; o, frontal aspect of head; p, left view of head. R. H. COBBEN: Eurasian Saldidae tribution to be added to those presently known. Eight of the new species described below be- long to a group of species which is treated here as the genus Macrosaldula. A preliminary dis- cussion on its status is given on page 232 and a key to the 21 species now recognized is added. I intend to devote a separate paper to the pressing problem of generic delimitations in the Saldidae after publication (Cobben in press) of descrip- tions of some twenty new taxa, mostly from Af- rica. I dedicate this paper to Prof. A. N. Kiritshen- ko (1884—1971), the founder of Russian saldid taxonomy who had already labelled some pres- ently described species as new to science in Rus- sian collections. coracina sp.n. body length in mm d 3.4—3.6 2 3.7—3.8 general coloration type series uniformly black, apex of wing often narrowly pale; pronotum entirely black, rarely with lateroventral pale streak structure of forewing strongly coreaceous and transversely vaulted; remnant of membrane not visibly demarcated from corium cuticle of forewing highly polished, entirely smooth without any sculpture, commissure between clavus and corium lined with a row of pits texture of dorsum hairless 217 DESCRIPTION OF SPECIES AND COMPARATIVE NOTES Halosalda coracina sp.n. (figs. 1 1-1; 2a, b). Description. — For measurements, see table 1. Typical Halosalda habitus, but dorsal cuticle completely bare, highly smooth and polished. Greece. The characters of this new species are pre- sented in comparison with conditions in the other two Halosalda species. Since only of H. lateralis (Fallen, 1807) and H. concolor (Pu- ton, 1880) macropters are known, comparisons below refer to semibrachypterous specimens (forewing more or less coleopterous due to sub- stantial reduction of membrane). concolor 3.2—3.7 3.6—4.0 variable (see figs. 102— 105 in Cobben, 1960); extension of black pigment spreading in eunomic series from mesocorium outward; predominantly dark specimens ın west mediterranean as coracina but less vaulted; membrane weakly set off from corium shiny, weakly rugose; suture between clavus and corium indistinct with very scattered short adpressed setae (distance between setae wider than length of setae) lateralis 3.24.1!) 3.6—4.6!) highly variable, but pale specimens predominating (see figs. 94—98 in Cobben, 1960); extension of dark pigment in wing starting from lateral sides not coleopteroid, dorsum in cross-section weakly convex; visible borderline between corium and membrane weakly shining, clearly rugose; suture between clavus and corium not obvious, but claval ridge along inner side of commissure distinct with rather dense regular coat of short decumbent setae (distance between setae much shorter than length of setae). 1) These numbers refer to specimens from various origins; the mean value for material from the west continental coasts is clearly higher than for specimens from the British Isles and countries bordering the Black Sea. 218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 ratio: width/length d 2.05—2.25 2.50—2.70 2.60—2.75 ef prenetum 9 2.35—2.50 2.65—2.90 2.70—2.95 scutellum anterior mid part wıth flat flat shallow depression head very polished, frons with rather shiny, frons rather dull, frons and vertex only few scattered setae; | somewhat rugose, thickly covered with pale - white parties of mouth moderately beset with setae (fig. 10); mouth sclerites mostly as in fig. short setae (fig. lm, n); sclerites extensively light- 11, sometimes as in fig. 1m pigmentation of mouth coloured; post-clypeus or in between fig. 1m and sclerites varying between __ transversely well- n, but lateral edges of fig. Im and n, and developed, fused with transverse band sometimes grading into mandibular plates and (postclypeus) never that fig. lo, particularly in lateral swellings between swollen specimens from Cyprus eye and antennal socket antennae dark brownish, inner side dark specimens: segment entirely pale, segment 1 and of segment 1 yellowish 1 and 2 pale, outer side 2 often with dark line along with brown streak; light- external side coloured specimens from Cyprus: entirely pale legs first acetabula largely dark specimens: first as in concolor pale; coxa blackish, femur acetabula narrowly or and tibia ochreous with largely pale; legs with dark longitudinal stripes short brownish stripes; lightest specimens: all acetabula and entire legs whitish genitalia parandria slender (fig. 2b) parandrıa blunt (fig. 2c) parandria blunt (fig. 2c) known geographic coast of NE Greece distribution Material. — Holotype (6), Porto Lagos, NE Greece, 23.viil.1983, leg. R. H. Cobben. Paratypes 9 3 4 9, idem. Holotype and paratypes in coll. Wage- ningen !), paratypes in Leningrad coll. and in coll. Drosopoulos, Athens. Comparative notes. — The type series, all strongly semibrachypterous, was collected on open patches of moist sandy soil in Salicornia vegetation in the dunes bordering the Aegean 1) The indication “coll. Wageningen” throughout this paper means: the collections of the Department of Entomology of the Agricultural University at Wa- geningen. mediterranean (Italy, France, Spain, Tunesia, Cyprus, Corfu) widely Palaearctic Sea. The species occurred simultaneously with H. lateralis. Since sampling was done in the eve- ning twilight and only two specimens of H. la- teralis were seen, the possibility of a difference in the daily period of activity between both spe- cies cannot be excluded. The most reliable char- acter of H. coracina separating it from the other two species, lies in the lack of any dorsal sculp- ture and pilosity. As regards the coloration, the new species forms the most dark extreme of a gradual series of colour morphs, in which A. la- | teralis represents the opposite extreme of pre- dominantly light-coloured specimens. H. con- color has a more or less intermediate position, at R. H. COBBEN: Eurasian Saldidae 219 Fig. 2. a—c, Halosalda. a, H. coracina, median endosomal sclerite; b, c, parandria of H. coracina (b), H. latera- lis and H. concolor (c). d—i, Saldula taiwanensis. d, frontal aspect of head of 3; e, paramere; f, penisfilum; g, left view of penis; h, parandria; i, left fore wing. j—l, Saldula inoana. j, parandria; k, paramere; |, frontal aspect of head of 3; m, Saldula uichancoi: frontal aspect of head of d. 220 Fig. 3. Saldula hasegawai, general facies of 9. least in the western mediterranean, where I of- ten collected it together with H. lateralis. population from Cyprus (near Akrotiri, 21.vi.1951, leg. G. Mavromoustakis), which on the basis of cuticular structure and paucity of setae definitely belongs to H. concolor, is con- siderably different. All its individuals are ex- tremely lightly pigmented. Some are even more pale than the lightest individuals of H. lateralis I TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 have seen, the mid frontal pale spots on the head conjugating with the transverse postclypeal swelling. This sample from Cyprus furthermore contains a high percentage of macropters (6 ma- cropteres, 4 semibrachypters), the first seen in H. concolor. Although Akrotiri is close to the | sea, the locality label mentions “in fresh water | marshes”. This fits in with my own and Josi- fov’s (1957) experience that the macropters of R. H. COBBEN: Eurasian Saldidae 221 H. lateralis are occasionally found outside the main habitat of the species. It is expected that H. coracina occurs further eastward in coastal salt-marshes in Turkey. My collections from more south-eastern and west- ern parts of Greece so far revealed only the presence of H. lateralis. The distribution pat- tern of H. concolor is still fragmentary, and col- lections containing material from southern ori- gins identified as H. lateralis need to be reinves- tigated. On my request, Dr Kerzhner checked the abundant material in the Leningrad Museum and concluded that it all refers to H. lateralis proper. This Euro-Siberian species occurs all over Europe inclusive of Scandinavia and its eastern range covers East-Mongolia and Trans- baical. Saldula hasegawai sp.n. (figs. la—g; 3) Description. — For measurements, see table 1. Rather small (3.0—3.6 mm), head and thorax black, glossy, wing dull-black with ochreous wing margin, semibrachypterous (membrane half-way reduced), whole body with long pilos- ity. Japan, Far East of the USSR. Head: black, spherical preocellar spot and all mouth-part sclerites, gular lobe and rostrum yellowish brown; glabrous yellowish white tu- mescence in between antennal socket and eye; dorsal surface with numerous erect dark setae as long as the trichobothrial setae and with recum- bent short golden setae; eyes with short setae; underside with semilong adpressed silvery setae. Thorax: black, glossy, densely covered dorsally with upstanding black setae and recumbent semilong light setae; pronotum with distinct collar, separated from callus by row of pits, lateral sides weakly convex or straight, frontal edges not wider than collar; dome about two- and-a-half times as long as posterior lobe, pos- terior border of dome lined with pits. Wing: ashy black, clavus without apical pale spot, endocorium entirely dark in specimens from Japan, with one slight spot in centre and another one on inner edge in specimens from E. USSR; exocorium convex laterally, with light brown lateral margin extending in light subbasal and apical spot; with long erect brown and re- cumbent, shaggy, semilong golden setae; mem- brane distinctly reduced, dark smoky with och- reous spots or entirely testaceous; hypocostal lamina without secondary oblique ridge; apex of hind wing reaching level of base of mem- brane. Extremities: ratio of antennal segments 1 : 2.1 : 1.5 : 1.5, segments 1 and 2 brown, shining, with erect brown setae, which are slightly lon- ger than diameter of segment, and some setae of greater length on segment 2; segments 3 and 4 dark-brown with short white setae and scat- tered dark erect bristles. Legs unicolorous light- brown, acetabula and coxae black; tibiae with erect brown setae, which along outer margin are longer than width of tibiae, and dark spines as long as tibial diameter. Other structures: rudiment of larval organ and sclerites of pregenital gland present; subge- nital plate of © broadly truncate (fig. 1f), grasp- ing plate of d with some 23 pegs (fig. 1b); par- andria and paramere as figured (figs. le, a, c), penisfilum coiled 212 times, endosomal sclerite of normal shape (fig. 1g). Holotype 6, length 3.16 mm, width 1.70 mm. Paratypes, 4 d, length 2.8—3.05 mm, width 1.6—1.7 mm; 59, length 3.2—3.7 mm, width 1.78—1.9 mm. Material. — Holotype (4), Japan, Osorezan, Ao- mori Pref., 2.viii.1953, leg. H. Hasegawa (in coll. Wa- geningen). Paratypes, 3 d 4 ®, idem (in Hasegawa coll.); 1 & 12, USSR, Sudzukhe (now Zapovednyy), 25 km S of Sokolovka Primorskiy Kray, coast of Japa- nese Sea, 22.viii.1959, leg. I. Kerzhner; 1 d, near Vla- divostok, 26.vii.1925, leg. Rostovykh (in coll. Lenin- grad Museum). Comparative notes. — Superficially this new species resembles on the one hand the Micra- canthia fennica group in general facies, pigmen- tation (fig. 4g—j) and shape of the paramere (fig. 4a—f), and, on the other hand, the Nearctic Saldula bouchervillei (Prov.) (fig. 4m), S. orbi- culata Uhl. (fig. 4n) and S. severini Harr., in colour pattern and pilosity. Particularly the re- semblance with S. orbiculata is rather strong, but unlike in that species the pruinose areas on the wings are lacking in S. hasegawai. Besides smaller specific differences (e.g. in shape of par- ameres, compare fig. la with figs. 4k, 1), S. ha- segawai deviates clearly from all these species in having a glabrous, lightish tumescence between antennal socket and eye, and in not having the secondary hypocostal ridge. Saldula taiwanensis sp.n. (fig. 2d—1i) Description. — For measurements, see table 1. Rather small (3.5—4.0 mm), short-haired, head and thorax black, moderately shiny, wings fully developed, with black, bluish pruinose 222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Fig. 4. a—j, Micracanthia. a—f, paramere; g—j, left fore wing. a, M. marginalis; b, M. humilis; c, M. drakei; d, M. husseyi; e, M. pumpila; f, M. floridana; g—i, M. marginalis; j, M. fennica; k, m, 0, S. bouchervillei. |, n, p, S. orbiculata. k, |, paramere; m, n, left fore wing; O, pP, pronotum. R. H. COBBEN: Eurasian Saldidae and white markings more or less like in the ge- nus Chartoscirta. Taiwan. Head: black, anteclypeus, maxillary and man- dibular plates largely yellowish, labrum black with central part lightish, apex of gular lobe yel- lowish, preocellar spot broadly touching eye margin (fig. 2d); short adpressed golden hairs, vertex with several semilong erect black setae, underside with dense silvery pubescence; ros- trum brown, reaching in between hind coxae. Thorax: black, rather shiny, with adpressed short golden pubescence, side margin of prono- tum straight or weakly concave, front edges slightly wider than collar; acetabula black. Wings with irregular golden pubescence, on mid part somewhat shaggy, clavus with only ap- ical, small lightish spot, embolium yellowish with black base; distribution of black, white and pruinose areas of corium and pigmentation of membrane as in fig. 21; hypocostal suture pre- sent. Extremities: antennae, length of segments given in table 1, nr. 3, segment 1 yellowish brown, underneath black; other segments dark brown with short brown hairs, additional erect hairs on segments 3 and 4 not longer than diameter of segment. Legs yellowish, coxae dark brown, femora ventrally with dark streak, flat sides with brown dots; knees and apex of last tarsal segment fuscous. Other structures: larval organ present, stig- mata not contacting side margins of sternites, male coupling plate with about 18 small pegs. Genital capsule of male with erect long brown setae on dorsolateral sides; parandria widely separated (fig. 2h); paramere with sharp pro- cessus hamatus (fig. 2e); length of penisfilum and phallic sclerites as in figs. 2f and g. Subgeni- tal plate of £ broadly rounded, whitish. Material. — Holotype (4), Taiwan, Baron-Nishi- mura, 10.viii.1941, leg. H. Hasegawa (in coll. Wage- ningen). Paratypes idem 8 d 3 @ (in coll. Hasegawa and Wageningen); M. Taiwan, Keishinryo, nr. Chu- chi, 1 d, 15.1v.1965, leg. T. Saigusa (in coll. Miyamo- to). Comparative notes. — The external aspect of S. taiwanensis resembles very much the Pal- aearctic Chartoscirta cincta (H.-S). The new species, however, lacks the leg-wing sound pro- ducing mechanism altogether; the unique shape of the plectrum is diagnostic for Chartoscirta. Other characters (shorter first antennal seg- ment, male genitalia) also prevent inclusion into Chartoscirta. The new species also has some su- perficial resemblance to species of the S. fletche- 225 ri group (e.g. S. fletcheri (Dist.), S. moana Dr., S. uichancoi Dr. & Viad.). Male genitalia (fig. 2), k), and other characteristics (fig. 21, m) of this group are quite different from S. taiwanensis. Saldula burmanica Lindskog, 1975 subsp.n.? (fig. 5a—d, f, g) Description. — For measurements, see table 1. Medium sized (3.44.6), mounticolous spe- cies of the orthochila group, coalblack with dense, often conspicuous semierect dark pubes- cence, shining head, thorax and lateral wing margin; wing mostly with some greyish white spots, clavus most often with subbasal and sub- apical spot; polymorphous, usually broadly subovate subbrachypterous. Himalayan moun- tain chain, India, Nepal. Head: rather shining, with recumbent light setae and some semilong dark setae which are shorter than the trichobothrial bristles, eye with scattered, very short setae; preocellar spot trian- gular, one side adjacent to eye; middle pair of trichobothrial setae on weak, black tumescence; anteclypeus with two brisles; mouthpart scle- rites pale yellowish, more darkened in female, gular lobe black, rostrum light or dark brown. Thorax: shiny, glossy, with some light ad- pressed setae and an irregular vestiture of erect short or semilong dark setae; pronotum (fig. 5g) with straight of slightly convex lateral sides, proximal side somewhat wider than collar, dome not reaching side margins of pronotum, central pit deep, posterior margin indicated with row of pits; first acetabula with narrow pale margin, acetabula 2 and 3 entirely black. Hemielytron (fig. 5f): dull, lateral explanate strip of exocorium and R + M ridge deep black, shining; other wing parts ashy grey and black with regular, rather short, semierect dark setae and some scattered golden adpressed setae, length of semierect setae subequal to width of hind tibia; clavus mostly with a small subbasal and subapical light spot, sometimes indistinct or absent; endocorium with varying number of small light spots, maximum number six, two near R + M ridge, four in distal part, light spots sometimes pruinose; exocorium with maximally three pale spots in the inner region and one larger spot distally near the wing margin; this latter spot remaining visible in otherwise en- tirely black specimens; membrane shining, dark-light pattern more or less as in fig. 5f, mostly subbrachypterous to varying degrees, sometimes tending to semibrachyptery at high altitudes (ca. 3000 m), hind wing as long as fore 224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Fig 5. a, Antenna of S. orthochila from W Europe (above) and from S. burmanica (below); b, median endosomal sclerite of S. orthochila (left) and S. burmanica (right); c, paramere of S. burmanica from Nepal (left), India (right); d, parandria of S. burmanica from India (above), Nepal (middle), of S. orthochila (W Europe); e, para- mere of S. orthochila from the Netherlands: f, g, S. burmanica; f, fore wing; g, pronotum; h, pronotum of speci- men of the burmanica group from Vietnam. R. H. CoBBEN: Eurasian Saldidae 225 wing or reduced till apex of clavus of fore wing; submacropterous condition of fore wing rare. Extremities: Antennae shining, segment 1 light brown with some semilong dark bristles; segment 2 light or dark brown with short dark setae and some erect semilong dark setae along median side, 2—3 times as long as width of seg- ment (fig. 5a below); segments 3 and 4 black; ratio of segments 1 : 2.5—2.6 : 1.7—1.8 : 1.9— 2.1. Legs shining, testaceous, with short light brown pilosity, coxae black, femora with dark blots on flat sides and ventrally with black stripe; base, mid part and apex of tibiae dark- ened, black spines as long as diameter of tibia, last tarsal segment dark brown. Other structures: larval organ absent, sclerite of pregenital gland present; subgenital plate of female broadly rounded, black, caudal margin sometimes narrowly pale; coupling plate of male with about 20 pegs in three rows, paran- dria broad, nearly adjacent (fig. 5d), paramere with acute-angled, black processus hamatus and pronounced processus sensualis with long, brown setae (fig. 5c); median endosomal sclerite as in fig. 5b, right; penisfilum coiled two times. Length of 10 d 3.44.1 mm, width 1.8—1.9 mm; of 5 2 3.94.9 mm, and 2.1—2.4 mm, respectively. Material. — India: W. Almora, 2 d 39, Kumaon, (no date), leg. H. G. Champion; Kumaon, Nainital, 1 2, (no date), leg. H. G. Champion; Chakrate Div. 2.300 m, 16, 1.vii.1932, leg. H. G. Champion; Versi gaon, Sallana Tehri, Garhwal, near stream, 1830 m, 1 3 1 2 2.vi.1946, leg. J. K. Uniyal; Uttar Pradesh, Mussoorie, 1500—2200 m, 19, 3—14.viii.1978 (Co- penhagen Zool. Mus. Exp.); Gopaldhara, Darjeeling, 1500—2000 m., 1 ®, 16.1x.1917, leg. H. Stevens; Sik- kim, Karponang 3.300 m, 2 2, 16—24.111.1917 (leg. H. S.); Sikkim-Nepal frontier, Tonglu, 3.350 m, 1 9, vii—viii. 1919, leg. H. Stevens; Nepal: R. G. Yack Exp. 1 6, 12.vii.1972, leg. R. A. Laurence; Pang- boche, 4000 m, shore of marsh, 14 d 4 2, 11.v11.1964; Junbesi, 2900 m, 1 ©, 28.vii.1964; between Those and Junbesi, 1 2, viii.1974, leg. C. Ravaccia; Puiyan, 2900 m, mist-forest, 19.vii.1964; Thangpoche, 3500 m, 1 6, 11.vii.1964; Pheriche, 4350 m, shore of swamp, 11 à 7 2, 10.vii.1964, all. leg. R. Remane; Alm Darghari, Maharigaon, 4000 m, 1 ®, ix.1971, leg. H. Franz; be- tween Mulkharka and Tare-Pati, 1 9, ix—x.1971, leg. H. Franz; Langtan vall., 1 6 1 2, 23.iv.1978, leg. H. Kraigher; Gufa, Terhathum Distr. 2950—3000 m, 1 d 1 2, 29.x.1979, leg. M. Tomokuni; Ting Sang La, 3400 m, 1 d 1 2, 13—15.1v.1973, leg. J. Martens. Comparative notes. — I must give an expla- nation for the description of and discussion on a taxon presented here, which was perfectly de- scribed by Lindskog (1975) as S. burmanica, based on 3 d and 4 ® from one locality (1200 m) in NE Burma. Since more than 15 years I had in my collection specimens from high alti- tudes in India which I described in manuscript as a new species. In recent years I saw addition- al material from many localities in the Hima- laya. Since my and also Lindskog’s (1975, p. 170) belief was that indeed a valid species was involved, I prepared its formal description for the present paper. The manuscript of the pre- sent paper was sent to Dr Lindskog for his com- ments. Meanwhile Lindskog had also received many additional specimens of “the new species” and he wrote me that he is now inclined to con- sider it at most a subspecies of S. burmanica Lindskog. Having seen now the holotype and paratypes of S. burmanica, which were kindly sent by Lindskog for comparison, I agree with Lindskog’s conclusion. Since the whole body of my manuscript had already achieved its final stage, inclusive the illustrations, I retain here my original text, with only some necessary amendments. I leave the final decision on the taxonomic status to Per Lindskog who is pre- paring a detailed study of the burmanica-ortho- chila complex. His paper will provide a much more detailed and geographical analysis than presented here. Saldula orthochila Fieber, the closest relative of S. burmanica, differs in the following re- spects: more slender (ratio body length: width about 2.1—2.2, as against 1.8—2.0 in the S. bur- manica form considered here); without long erect black setae on head and thorax and with- out erect dark pilosity on hemielytra; recum- bent light setae more numerous; head and tho- rax less shining; first acetabula broadly light; lateral strip of fore wing dull, membrane for the major part hyaline (see for more differences Lindskog, 1975, p. 165). S. orthochila has a very wide distribution covering nearly all West Eu- ropean countries, and it is recorded eastwards from S Russia, Turkestan and Siberia (Cobben, 1960). Iran is added here as a new country re- cord (Mazanderan, Chalus-valley, 1300 m, 1 9, 13.viii.1968, leg. Heinz). I have seen material from 3000m altitude in Tibet (Supi River, no date, 6 d 14 2, leg. H. G. Champion, in Coll. BMNH, London) and from Kashmir (Rukshu, Tso-Morari, near frontier Tibet, 1 d, vii.1914, leg. G. Babault, in Coll. Smiths. Inst. Washing- ton). These match specimens from W Europe, except for varying colour of second antennal segment (dark brown to light brown), for some long erect setae on head and pronotum, and for 226 somewhat denser and longer pilosity on the wings. Increasing pilosity at higher altitudes is also seen in other saldids (e.g. Saldula saltato- ria), but the pilosity of S. orthochila remains much less dense than in S. burmanica. The semibrachypterous orthochila from Tibet have further reduced hind wings, reaching about the apex of the clavus; the extending setae on the second antennal segment are of varying length, mostly shorter than in western populations. On the basis of all these characters, the orthochila material from Tibet seems somewhat interme- diate between typical S. orthochila and S. bur- manica. A study of the geographic variation of additional populations from western and more eastern parts of the Himalaya is needed in order to define subpopulations of S. orthochila and S. burmanica. To my surprise, I came across one female from N Vietnam, which means a considerable extension of the known orthochila-burmanica chain. The specimen in the Smithsonian Institu- tion, Washington, is labelled: Tonkin, Chapa, v.1916, leg. R. V. de Salvaza. It is long-winged, 4.51 x 2.20 mm, and conforms most to the de- scription of S. burmanica (second antennal seg- ment light-brown with mediolateral setae about two times as long as width of segment; area around subapical spot of clavus concolorous with the rest of clavus). The aspect of the pro- notum (fig. 5h) seems to differ somewhat in that it tapers more narrowly towards the collar than in orthochila and our form of burmanica (fig. 5g). The taxonomic evaluation of the Vietnam individual must wait till males are available from that territory. It presumably lives there at high elevations. An extensive survey of the habitat and ecology of the orthochila group of species 1s presented by Lindskog (1975). Another species of this group, defined by the character set: sec- ond antennal segment with some erect bristle- like setae and absence of the larval organ, is $. nobilis Horv. with a Central European-Asiat- ic distribution. It is a mountainous species and the known records are scattered (see map 1). I have now seen the first specimens from Japan (Hokkaido, Kiyokawa near Ashora, 2 d 19, 7.v11.1958, leg. S. Miyamoto; Berabonai, Asho- ro, 1 2, 8.v11.1958, leg. S. Miyamoto; Kuttyaro- ko, 1 2, 11.viii.1937, leg. S. Asahina) and from China (Manchuria, 1 ©, 25.vii.1943, leg. E. Ka- wase). Lindskog (1975) referred to S. boucher- ville: Prov. (= S. illinoiensis Dr.), a Nearctic species exhibiting phenetic affinities to the or- thochila group. This species indeed has no larval TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 organ as is typical for the orthochila group. A reliable cladistic analysis of the species clusters of the genus Saldula would ultimately bring out whether the Palaearctic and the Nearctic assem- blage of species, lacking the larval abdominal organ (Cobben, 1957, 1959), form together a monophyletic group. An independent loss of that organ seems probable. Other species in the Nearctic Region, which lack such organ, belong to different complexes. They are S. villosa Hodgd. (California, Oregon) on the one hand, and S. laevis Champ. and S. sulcicollis Champ. both from Guatemala and Mexico, on the other hand. Oiosalda caboti Drake & Hoberlandt, 1952 (Colombia) seems to be most closely re- lated to the latter two species, but actually pos- sesses a larval organ. Definitely not closely re- lated to the orthochila group, but nevertheless without a larval organ, is a complex of species . from India and surrounding countries (S. cham- pioni Dr., edita Dr. & Hoberl., fletcheri Dist., pusana Dist.). Saldula sibiricola sp.n. (fig. 6f—n) Description. — For measurements, see table 1. Medium-sized (4.3—5.7 mm), belonging to the orthochila group, very close to S. nobilis Horvath, but with more extended testaceous elytral markings. Siberia. Head: wide (0.7—0.8 x width of pronotum), shiny black, preocellar spot elongate triangular with longest side along eye; all mouth sclerites lightish in à, black in © except for anteclypeus and labrum, which are sometimes fuscous; with shaggy adpressed lightish setae; postclypeus, frons and vertex also with many extending dark setae nearly as long as trichobothria; eyes with scattered short setae; rostrum dark brownish, reaching or slightly surpassing hind coxae. Tho- rax: black, shiny, with irregular golden pubes- cence and erect brown setae; pronotum narrow (ratio length/width 0.5 in submacropterous, 0.35 in macropterous specimens), lateral sides weakly concave, fore edges not much wider than the distinct collar; dome well-developed and elevated, reaching lateral sides, length me- dially 1.45 X posterior part of pronotum; first acetabula entirely white, 2 and 3 with pale apex. Wings: mostly submacropterous fore wings, hind wings reaching middle of membrane, one female macropterous, weakly shiny, with scat- tered golden recumbent setae on corium and veins of membrane; corium and clavus with dense pilosity of erect long brown setae (about R. H. COBBEN: Eurasian Saldidae 207 Fig. 6. a—e, Macrosaldula from Japan. a, M. miyamotoi, profile of pronotum and scutellum; b, idem of M. shi- kokuana; c, M. miyamotoi, paramere; d, e, hind tibia and antenna of M. shikokuana, respectively. in, Saldula sibiricola. f, paramere; g, apex of paramere; h, grasping plate of d; i, median endosomal sclerite; j, paramere; k, base of penisfilum; |, pigment variation of fore wing; m, outline of subgenital plate of 2; n, gynatrium and sper- matheca; o, S. nobilis, paramere. 228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 0.2 mm long); ground colour testaceous, dark pigment brownish (variation, fig. 61), three of the five specimens have a light proximal spot on the clavus; oblique hypocostal ridge present; membrane entirely light-coloured, basally with weak brown shade in dark specimens, veins light brownish. Extremities: first two antennal segments shi- ny, yellowish, base of 1 sometimes darkened underneath, 3 and 4 black; segment 2 with obliquely extending brown pilosity, along the distal median side somewhat longer than diame- ter of segment, and with two dark, long setae medially at the middle (about 4 X longer than diameter of segment). Legs inclusive of coxae yellowish; flat sides of femora with some fus- cous spots, apex of tibiae and last tarsal segment | infuscate; short lightish pubescence, dark spines of third tibiae as long as diameter of tibia. Other structures: abdomen dark brown, apex of sternites very narrowly pale; stigmata close to lateral margin of sternites; larval organ ab- sent; distal prolongation of subgenital plate of female semicircular, whitish; male grasping plate, parandria, paramere and penisfilum as fig- ured (fig. 6f—k, m, n). Holotype (d), length 4.3 mm, width 2.1 mm. Length and width of 3 submacropterous ® varying from 4.5—5.7 and 2.3—2.7 mm, re- spectively; of 1 macropterous ® 5.15 and 2.2 mm. Material. — Holotype (4), USSR, Transbaicalia, Kokuy, about 180 km ENE of railway station Urulga, 18.v1.1909, leg. A. Keller (in coll. Leningrad). Para- types: idem 1 d 1 © (in coll. Wageningen); mountains SW of railway station Koktuma, Dzhungarski Alatau Mts, Kazakhstan, forest gorge, 1 macropterous ©, 26.v1.1962, leg. G. Medvedev; 25 km ESE of An- dreevka, Dzhungarian Alatau Mts, Kazakhstan, in a narrow forest along river, 1 @, 7.vii.1978, leg. I. Kerzhner; Otar, region of Verny (now Alma-Ata), 1 2, 29.v1.1922, leg. A. Reichardt; Temirtau, near Kara- ganda, 1 ©, 11.vu.1961, leg. Asanova; river Tentek near Ursh-Aral, 1 d 1 2, 9.vii.1978, leg. I. Kerzhner. Comparative notes. — There are no clear-cut morphometric differences between S. sibiricola and S. nobilis (the paramere of S. sibiricola could be studied in only one ¢ available). The wing patterns are, however, so strikingly different that I am convinced, from my experience with other sibling complexes revealing fewer differ- ences (Cobben, 1960, 1961, 1980a, b), that in the present case we are confronted with two re- productively isolated populations. I have seen typical S. nobilis from Germany, Csechoslova- kia, USSR, China and Japan. The material seen from Asia is as following (see map 1). USSR: Verkhnyaya Mol’ka, Balagansk steppe, Irkutsk region, 1 6, 21.vin.1931, leg. Zakhvatkin; Igir- ma, Irkutsk region, 1 ?, 17.vin.1966, leg. Zhe- rikhin; Khabarovsk, 1 ©, 18.vu.1931, leg. V. Pereleshina; Khabarovsk-Ussuri region, 1 6, lEva 1977 leg RESET rica; Aid en OS CE 3.vii.1977, leg. Stys & Davidova; idem 1 9, 8:v11.1977, leg. Kr Hürka; idem 37821335 1.vii.1978, leg. Stys & Vilimovä; Mongolia, Batsiret, 1 9, 7.v111.1974, leg. A. Seifert. China: Manchuria, 1 9, 25.vii.1943, leg. E. Kawase. I added on map-1 one locality in the Amur region and one in China (Kuku-nor lake); this material in the Leningrad collection has been checked by dr. Kerzhner (in litt.). Japan (new country re- cord, formerly published as S. scotica Curt. by. Hiura, 1967): Hokkaido, Kiyokawa near Asho- ro, 2 d 2 9, 7.v11.1958, leg. S. Miyamoto; Kut- tyaroko, 1 @, 11.viu.1937, leg. S. Asahina; Bek- kai-mura, 1 ®, 2.vin.1967, leg. T. Saigusa; Ata- ruma-dake, 1 2, 11.vii1.1967, leg. A. Nakanishi. Most of the material is semibrachypterous, some are submacropterous. The question of the status of S. reuteri Jak. must be considered here briefly. Jakovlev described this species from Sı- beria in 1889 without reference to S. nobilis de- scribed by Horvath five years earlier. Part of the type series of S. reuteri, available to me by the courtesy of Dr Kerzhner, conforms exactly with S. nobilis. S. reuteri was treated as a variety or a synonym of S. nobilis by most subsequent authors such as Drake & Hoberlandt (1951), Lindskog (1975), Vinokurov (1979a, b , c), but listed again as a species propria by Hoberlandt (1971b) from Mongolia without further com- ments. All specimens of S. nobilis have a uniform, contrasting black-white wing pattern. The proximal basis of the clavus is black; rarely there is a very small white spot in the edge bor- dering the corium (just the opposite side in S. s7- biricola, fig. 61). The distal large white spot of the exocorium, which persists in all specimens of S. nobilis, presents a striking resemblance to Teloleuca pellucens. In contrast, the light colour of the corium of S. sibiricola is testaceous and the brown pigmentation is only vaguely indi- cated (fig. 61). I dare to predict that specimens with darker patterns than drawn in fig. 61, right, will eventually be found, and that their distal endocorial spot will be much more reduced than in S. nobilis. R. H. COBBEN: Eurasian Saldidae 229 The conclusion that sibiricola is a valid spe- cies and not just an ecotype, is further strength- ened by the fact that the eight specimens origi- nate from five different localities whithin an area of roughly 1200 km? between 70—95° longit., 55—40° latit. (see map 1). Its range is more or less surrounded by that of S. nobilis which extends from western Europe to eastern USSR and Japan. Although this distribution pattern suggests vicariance between both spe- cies, Dr Kerzhner wrote me that S. sibiricola ap- parently is a lowland species, in contrast to S. nobilis. All records of the new species origi- nate from steppe or even semidesert regions outside the true mountainous regions, mostly at altitudes between 100 and 400 m. S. sibiricola might be more thermophilous than S. nobilis which prefers damp situations at highter alti- tudes. Macrosaldula clavalis sp.n. (fig. 7a—e) Description. — For measurements, see table 1. Moderate size (4—5 mm), without erect long setae, pronotum with pale side margins, clavus with lightish basal stripe, wings extensively marked with light pattern, close resemblance to M. kaszabi (Hoberlandt, 1971). USSR. M. clavalis sp.n. and M. kaszabi (Hober- landt) can be distinguished as follows. clavalis kaszabi length of antennae in relation to 22-23 X 2x width of head ratio length antennal segments 3 + 1.25—1.4 11.2 4 to that of 2 ratio pronotum width/length 3.032 27 head and pronotum shiny dull pubescence short, not dense, golden rather dense, silvery mouthpart sclerites of ? predominantly dark, only lightish mandibular plate and apex of anteclypeus lightish (fig. 7d) wing margin partly dark entirely lightish inner base of clavus 7a) tibiae ring Material. — Holotype (4), USSR, Transcaucasia, Tshakvis-tavi, Adzharia, 15—20 km NE of Batumi, 21.v11.1949, leg. Kiritshenko (in Leningrad coll.). Par- atype d, idem (in Leningrad coll.), 1 ©, idem (in coll. Wageningen). Comparative notes. — Both the new species and M. kaszabi (Hoberl.) share a pale dot at the origin of the middle cephalic trichobothria, pale acetabula and pale pronotal side margins. The gular plate of M. clavalis 8 is of light colour. The paramere of M. clavalis has a longer pro- cessus hamatus (fig. 7b, e) than in M. kaszabi as figured by Hoberlandt (1971 b, figs. 10, 11). The type locality of the new species in Georgia is about 4500 km west of the range of M. kasza- bi in Mongolia. The differences with other Ma- crosaldula appear from the key to Macrosaldula with longitudinal pale stripe (fig. dark coloured except for subapical without basal stripe lightish except for dark base and apex species presented below. Comments on the ge- neric status of Macrosaldula are presented fol- lowing the description of the next species, and on page 254. Macrosaldula inornata sp.n. (figs. 8, 9 i—n) Description. — For measurements, see table 1. Medium-sized (4.3—5 mm), slender, full- winged, short-haired, predominantly straw-yel- lowish, facies superficially Pentacora-like (fig. 8). Iraq. Head: black, weakly shining, with recumbent silvery hairs; postclypeus, frons and vertex with ocelli flat; transverse swelling broadly devel- oped above insertion of antenna; preocellar and frontolateral spots, border of upper notch of eye, transverse swelling, anteclypeus (except 230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Fig. 7. a—e, Macrosaldula clavalis. a, corium and clavus of left fore wing; b, apex of paramere; c, male grasping plate; d, front view of head; e, paramere. f—k, Macrosaldula koktshetavica. f, paramere; g, base of penisfilum; h, median endosomal sclerite; i, rudiment of larval organ; j, parandria; k, pigment variation of corium and clavus. 1, Macrosaldula oblonga, pigment variation of wing. R. H. COBBEN: Eurasian Saldidae 231 Fig. 8. Macrosaldula inornata, holotype 4. base), maxillary plate, gula and labrum yellow- ish; first visible segment of rostrum light, re- mainder dark brown, extending to middle cox- ae. Thorax: nearly dull, densely covered with recumbent golden pilosity dorsally and medi- um-long silvery hairs ventrally; pronotum tra- pezoid, with pale lateral margins, dome rather flat; acetabula light. Wings: dull, with rather regular short, silvery pubescence, corium and clavus nearly unicolo- rous straw-yellowish, membrane slightly smo- ky; eight specimens have the lightest pattern as shown in fig. 8; in two males the dark design on the corium is only little enlarged, but the pale clavus tends to be broken up by pigment trans- versally in the middle; hypocostal lamina nar- row, without oblique ridge; area of wing margin of female serving for attachment of male grasp- ing plate nearly imperceptibly differentiated; hind wing nearly as long as forewing. Extremities: antennae slender, only with very short hairs (except for the erect ones on 3rd and 4th segment); segment 1 stout, yellowish, 2 light brown, 3 and 4 dark brown; legs straw- yellowish, with brown patches as in fig. 8, coxae black with light apex; legs with very short sil- very pubescence, spines on last tibia brown, not longer than diameter of tibia; tibia 3 weakly curved inwards. Other structures: abdomen dark brown, dis- tal margin of sternites light; rudiment of larval organ and sclerite of pregenital gland present; stigmata close to but not touching side margins of sternites; subgenital plate of female with black base and white, truncate distal prolonga- tion (fig. 9k); ovipositor with eight teeth (fig. 232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 91); spermatheca asymmetrical, piriform; sper- mathecal duct gradually expanding and opening into spermatheca without a flange (fig. 9n). Male grasping plate with some 25 semi-long pegs; male genital structures as depicted in fig. 9, j,m Holotype 6, length 4.3 mm, width 1.7 mm. Length and width of 4 & 4.3—4.5 mm and 1.7—1.8 mm, of 5 ® 4.7—5.0 mm and 2.0—2.1 mm. Material. — Holotype (4), Iraq (Sept.), Prov. Mo- sul, near Agra, Salta-ravin, 24.vi.1958, leg. R. Re- mane. Paratypes 4 d 5 ©, idem. Holotype and para- types in coll. Wageningen, paratypes in Remane coll., Marburg, BRD. The species was collected reach with M. variabilis variabilis (H.-S) in the same habitat: stones in and along mountain-river (altitude between 600 and 1000 m). Comparative notes. — Reuter (1895, 1912) was the first to recognize a scotica species-group within Acanthia (= Saldula), which possibly might deserve the status of subgenus. He ın- cluded in this group S. jakovleffi, oblonga, rivu- laria, scotica, variabilis and, with some reserva- tion, koreana and mongolica not seen by him. To this group I added a new species from Spain (Cobben, 1959), characterizing the species- group by stout body dimensions, proportional- ly long antennae and the same type of median sclerotized structure of the penis. The taxon Macrosaldula was first informally introduced as a subgenus by Southwood & Leston (1959) in order to separate S. scotica (Curtis, 1833) from typical Saldula species, and formalized as such some years later (Leston & Southwood, 1964). The subgenus Macrosaldula subsequently was given generic rank by Wroblewski (1968) and Polhemus (1977). However, the antennal ratios given by Leston & Southwood to distinguish Macrosaldula from Saldula species are not at all exclusive, and certainly they are not exclusive from other genera. Awaiting a critical analysis of generic groupings (Cobben, in prep.), I pro- visionally follow the usage of recognizing a Ma- crosaldula clade, inclusive of the lapidicolous M. inornata described above. Although aberrant in the paucity of dark pigment, it shares with other Macrosaldula species the shape of the median endosomal sclerite (fig. 9j) and has the following plesiomorphous conditions in common: male grasping plate with oblong pegs and hairs, ab- sence of the secondary ridge on the hypocostal lamina and absence of the spermathecal flange. Lindskog (1975) suggested that also the Nearc- tic Saldula andrei Drake (= S. azteca Drake & Hottes) and S. nigrita Parshley belong in the Macrosaldula group, but these species do not reveal the combination of characters given above. Macrosaldula kerzhneri sp. n. (fig. 10a, b, d—i, n) Description. — For measurements, see table 1. Stout (5.9-—7.2 mm), rather dull, erect semi- long pubescence, predominantly black with nar- row testaceous seam along lateral wing margin (fig. 101, n). USSR, Kazakhstan. Since the general facies resembles a number of other dark-coloured species, only some main characteristics are mentioned here. The testa- ceous wing margin (fig. 101, n) separates this species from all other congeners. The semilong setae (length subequal to diameter of hind tibia) on the wings and scutellum are dense, on head and thorax sparse. Dorsum in addition with a regular coat of short decumbent silvery hairs. All mouth sclerites entirely (4) of partly (9) yellowish. Antennae and legs predominantly blackish or dark brown, light-dark pattern as in most other species; first antennal segment of 4 testaceous on inner side. First acetabula entirely and second and third apically yellowish. Male genital structures as drawn in fig. 10a, e—h. Male grasping plate with about 20 elongate pegs and some stiff setae medially (fig. 10b). Material. — Holotype(d), S. Kazakhstan, 20 km N of Kentau, Karatau Mts, 27.v.1966, leg. Arnoldi (in coll. Leningrad). Paraypes, 2 ©, idem, 26.v.1966, leg. Kerzhner; 1 ©, 24.v.1966, leg. Gurjeva; 1 2, Atshi- say, river Teresakan, Karatau Mts, 31.v.1936, leg. Lu- kyanovitsh. For the location of sampling sites, see map 2. Comparative notes. — The wing pattern with the neatly parallel ochreous costal margin and the pale first acetabula separates M. kerzhneri from other species with a more or less dense pu- bescence. To these belong M. scotica, madonica and tadzhika in which, moreover, the setosity extends laterally beyond the pronotal margin. The paramere without distinct processus sen- sualis and slender, sharp processus hamatus (fig. 10e, f) and the male grasping plate with two rows of pegs (fig. 10b) differs from those in M. scotica (fig. 10j—m and 10c). Further differ- ences between M. kerzhneri and all other re- lated species can be extracted from the key to Macrosaldula species. R. H. COBBEN: Eurasian Saldidae 233 7 rk 4 PONT AW TY (TICO VA 025 Fig. 9. Macrosaldula species. a—h, M. koreana; a, b, paramere; c, plane and side view of median endosomal sclerite; d, grasping plate of d ; e, base of penisfilum; f, parandria; g, apex of paramere; h, sawing blade of ovipo- sitor. i—n, M. inornata. i, paramere, plane side and inner view (right); j, median endosomal sclerite; k, female subgenital plate; 1, sawing blade of ovipositor; m, parandria; n, gynatrium with ring gland and spermatheca. 234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Macrosaldula koktshetavica sp.n. (fig. 7f—k) Description. — For measurements, see table 1. Medium-sized (4.3—5.5 mm), full-winged, weakly shiny, yellowish brown, exocorium, legs and acetabula predominantly light-coloured, with short adpressed and semilong erect pubes- cence. USSR, Kazakhstan. - Head: black, weakly shiny, sparce silvery pu- bescence, several erect dark setae on frons and vertex in addition to the six trichobothria; preo- cellar spot, mouth sclerites and gula yellowish in both sexes; rostrum dark-brown, except for first short visible segment, extending in between middle coxae. Thorax: black, shiny, with rather dense silvery or golden pubescence and scat- tered semi-erect, semilong setae; first acetabula broadly and other acetabula narrowly margined with pale colour. Wings: weakly shiny, pubescence as on tho- rax; colour pattern not very contrasting, euno- my as in fig. 7k (based on 38 specimens); the spreading of dark pigment starts from the exo- corium, but the edge of the outer wing margin is always narrowly brown even in the pale speci- mens; in the lightest extreme the inner base of the clavus bears a narrow pale spot; the dark ex- treme approaches the general wing pattern of M. oblonga (fig. 71); membrane for the major part light-greyish, also in darkest specimens, veins light-brown; hypocostal lamina narrow, without oblique alae Extremities: first segment of antennae black ventrally, yellowish dorsally, other segments dark brown, the very base of second lightish; pubescence very short, all segments with some erect semilong dark setae. Legs predominantly yellowish, underside of femora dark brown, flat sides with some fuscous spots, apex of tibiae and last tarsal segment brownish, pubescence very short, dark spines of hind tibia almost as long as diameter of tibia. Other structures: abdomen brown, distal margin of sternites lightish; distal prolongation of subgenital plate of female truncate, white. Male grasping plate with short pegs and spinous setae in the median edge. Base of penisfilum with nearly two coils (fig. 7g). Parandria, para- mere, and endosomal sclerite as in fig. 7}, f, h. Holotype (4), length 4.3 mm, width 1.9 mm. Length and width of d varying from 4.3—4.7 and 1.8—1.9 mm, respectively; of 9 from 5.0— 5.5. and 2.1—2.3 mm, respectively. Material — USSR, holotype (4), Borovoe, Koktshetav region, Kazakhstan, 27.vi.1932, leg. V. Popov (in coll. Leningrad). Paratypes: 5 d 17 9, idem; between Stshutshinsk and Barmashi, Koktshe- tav region 8 d 99, 4 larvae, 23.vi. and 1—2.vii.1982, leg. Filipyev (in coll. Leningrad, coll. Wageningen). Additional 50 specimens, not seen by me, from both localities are in the Leningrad Museum. Dr Kerzhner informs me that the Koktshetav Hills form an isolated mountain-massive in North Kazakhstan, surrounded by steppes and covered by birch and pine forests. Comparative notes (see key to Macrosaldula species). Macrosaldula miyamotoi sp. n. (fig. 6a, c; 14 f) Description. — For measurements, see table 1. Medium-sized (4.4—5.7 mm), slender, macropterous, predominantly black with short . and semilong pilosity, corıum mostly with rath- er contrasting yellowish markings (fig. 14f), membrane dark smoky. Japan. Head: black, with erect black setae somewhat shorter than trichobothrial setae; lightish preo- cellar spots large, broadly touching eye and nearly extending to ocellus; mouthpart sclerites yellowish in male, base of anteclypeus, median part of transverse swelling and margins of la- brum darkish, in female maxillary plate in addi- tion black; gula black; rostrum dark brownish, reaching hind coxae. Thorax: shiny black with sparse short adpressed golden pubescence, in addition dorsally with erect semilong and a few long dark setae (fig. 6a), which extend from straight lateral margin of pronotum; acetabula black, rarely with light margin. Wings: scattered short golden setae, numer- ous dark erect semilong setae with curved apex; eunomy of weakly shining forewings as de- picted (fig. 14f); yellowish spots rather con- trasting with black ground-colour (less in speci- mens of Ohshimizu), dark area of distolateral part of endocorium deeply black; pale stripe in proximal base of clavus rarely present; mem- brane predominantly fuscous also in light-col- oured individuals. Extremities: Antennal segments dark, first segment medially with pale line, 2 with short se- tae. Legs predominantly dark-coloured, pubes- cence short; trochanters, upper and underside of femora and often subapical ring of tibiae and second tarsal segment lightish. Other structures: caudal extension of female subgenital plate truncate, white; paramere as in fig. 6c. R. H. COBBEN: Eurasian Saldidae 235 Fig. 10. Macrosaldula. a, b, d—i, n, M. kerzhneri. a, base of penisfilum; b, grasping plate; d, parandria; e, f, paramere, f, viewed in direction of arrow in fig. e; g, h, median endosomal sclerite, plane and lateral view, re- spectively; i, n, left fore wing. c, j—m, M. scotica. c, grasping plate; j—m, parameres of specimens from the Netherlands (j, 1) and Austria (m). 236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Holotype (6), length 4.9, width 1.9 mm. Length and width of 12 d varying from 4.4— 5.0 mm and 1.9—2.0 mm, respectively; of 8 ©: 5.3—5.7 and 2.3—2.5 mm. Material. — Holotype (d), Japan, Honshu, Nagano Pref., Kamikochi, 10.1x.1951, leg. H. Hasegawa (in coll. Wageningen). Paratypes: Idem 1 2; Honshu: Nagano Bat, Meck, SAIS MSC 25 a= 14.viii.1978, leg. M. Satô; Gifu Pref. Oppara, 9 d 39, 15:vi.1978, leg. M. Sato; Meoto waterfall, Takasu Vil- lage, 12 & 3 Q, 11.viii.1980, leg. J. T. Polhemus & M. Sato; Okumino, Hirugano Heights, 5 d 1 8, 12.vii.1980, leg. J. T. Polhemus & M. Sato; Izu Pen- insula, Nikai-daru Falls, 5 d 3 9, 29.1x.—3.x.1980, leg. M. Tomokuni; Izu Peninsula, Yagashima-cho, Kanogawa River, 4 & 5 2, 3.x.1980, leg. M. Tomoku- ni; Miyagi Pref., Futakuchi, Natori-gun, 1 9, 20.viii.1977, leg. M. Tomokuni, Toogatta, 1 ®, 19.viii.1977, leg. M. Tomokuni; Shosenkyo Kai, 1 d 1 2, 11.viii.1959, leg. S. Miyamoto; Yamanashi Pref., Masutomi, 1 d, 25.vu.1963, leg. T. Saigusa; Gumma Pref., Oze, 3 d 2 2, 5.ix.1952, leg. H. Hasegawa; Ohshimizu, Oku-Nikko, 3 d 49, 18.vi1.1940, leg. M. Hanano; Kyushu: Chikushi-Yabakei near Fukuoka, 2 3, 30.ix.1956, leg. S. Miyamoto; Ino near Fukuoka, 1 3 2 Q, 14.vii.1965, leg. S. Miyamoto; Fukushima, Chikugo, 1 d, 26.viii.1952, leg. S. Miyamoto; Fuka- goya, Chinugo, 1 ©, 6.vii.1952, leg. S. Miyamoto; Ya- kushima, Miyanoura, 1 d, 29.viii.1953, leg. Takeya & Hirashima. (Paratypes in coll. of Hasegawa, Miyamo- to, Polhemus, Jap. Nat. Museum, Kyushu Univ. and in coll. Wageningen). Comparative notes. — This species is appar- ently widely distributed along stony banks of rivers on the southern main islands of Japan. The lightest coloured specimen (fig. 14f) comes from Yakushima, a small island close to the southern coast of Kyushu. Colour-interme- diates with dark coloured specimens occur without any morphological differences on Honshu. M. miyamotoi and the related M. shi- kokuana from Japan, described below, are close to continental Macrosaldula species of the ob- longa group. Their eunomy, however, 1s marked by the absence of the light spot on the proximal part of the endocorium, even in the palest specimens (compare fig. 14f with figs. 71, 14d, e). Macrosaldula oblonga acetabularis subsp. n. (fig. 12f) Separable from the nominate form oblonga Stal, 1858, by the entirely whitish first acetabula (always black in the long series of typical ob- longa which I have seen). In two of the four specimens of the new subspecies the second ac- etabula have a pale margin. The ratio length an- tenna/width pronotum in two males of subsp. acetabularis is 1.64—1.74, and in 10 males of subsp. oblonga 1.85—1.90. These ratios could not be checked for females, since the single fe- male of acetabularis has incomplete antennae. The parameres of two males of the new subspe- cies have a straight processus hamatus (fig. 12f), whereas this is slightly curved upwards in subsp. oblonga (fig. 12a). Material. — 3 d 1 @, Zaisan, river Karasu, E. Kazakhstan 20.vi.1965, leg. J. Sukatsheva (holotype d in Leningrad coll., others in Popov coll. (Moscow), coll. Wageningen). This subspecies was collected si- multaneously with M. jakovleffi Reut. Comparative notes. — The locality of this subspecies is about 500 km outside the presently known range of the nominate form (see map 2). . The distribution pattern of M. oblonga oblonga covers the mountainous regions of Mongolia and Transbaical between 90 and 118 degrees of longitude, that is about 2000 km. The altitude of the type locality of M. oblonga acetabularis west of this range is not mentioned, but it may be lower than where the nominate form occurs. According to Hoberlandt (1971b), the occur- rence of the nominate subspecies in Mongolia is between 1200 and 2100 m. Macrosaldula shikokuana sp. n. (fig. 6b, d, e) Very similar to M. miyamotoi in dimensions (table 1) and coloration, differing only in the much longer pilosity. Second antennal segment with a few erect semilong setae along the medi- an side (fig. 6e). Erect hair-dress on dorsal side more conspicuous and somewhat longer than in previous species (fig. 6b), laterally extending over a distance of 0.23 mm beyond the margin of the pronotum. Tibiae with silvery adpressed pubescence and erect brown setae which are longer than the diameter of the tibia; the distri- bution and length of these curled setae on the hind tibiae are indicated in fig. 6d. The male paramere is much like in M. miyamotoi (fig. 6c). The wing patterns of the holotype and paratype are more or less like fig. 14f, right. Holotype (6), Japan, Shikoku, Omogo, Iyo, 14.vii.1952, leg. T. Ishjara & S. Miyamoto. Paratypes: idem, 3 d 3 ® (holotype and paratypes in Jap. Na- tional Museum, Tokyo; paratype in Polhemus coll., Englewood). R. H. Cossen: Eurasian Saldidae 237 Fig. 11. Macrosaldula species. a, M. simulans, parandria; b, M. rivularia from Mongolia, parandria; c, idem, left, median endosomal sclerite, plane side; right, left view of median and lateral endosomal sclerites; d—g, para- meres; d, M. monae; e, f, M. rivularia, Mongolia; h, apex of paramere of M. rivularia from Alaska; i—k, M. violacea; i, parandria; j, median endosomal sclerite; k, side view to show the placement of stigmata in between sternites and tergites. 238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Macrosaldula simulans sp. n. (fig. 11a) Description. — For measurements, see table 1. Stout (6.3—6.9 mm), unicolorous dullish black species with short silvery recumbent se- tae; very close to M. rivularia, Sahlb. East- USSR. The most reliable distinguishing characters with M. rivularia are as follows: middle pair of cephalic trichobothrial setae not originating from light spot (such a spot is present in all specimens of M. rivularia I have seen from Si- beria and Alaska, and also in the Alaskan M. monae Drake), and proportionally longer an- tennae and legs. The ratio antennal length/head width is 2.73—3.06 in M. simulans (n = 8) and 2.12—2.4 in M. rivularıa (n = 8). The ratio length of third tibia/head width amounts to 2.55—3.03 and 1.91—2.21, respectively. These ratios could be influenced by allometry of the appendages because the new species is on the average larger (6.3—6.9 mm) than M. rivularia (4.46.5 mm). However, the discrepancies of the ratios between the two species remain as striking in specimens of equal size. M. simulans differs further weakly in having a somewhat more dull pronotum, dark anteclypeus (also in the male), inner side of first antennal segment weakly lightish in male, entirely dark in female. Genital structures are similar to M. rivularia, but the parandria of the holotype of M. simulans reveals a small median process (fig. 11a), which lacks in M. rivularia (fig. 11b). Additional material is needed to check whether this is a constant feature of difference. Holotype (4), USSR, river Buren, near Ilyinka (Bulun-aksy), (Tuva region), 14.vi.1949, leg. Per- evoztshikova, in Leningrad Museum. Paratypes: USSR: 1 ®, Krasnoyarsk, 3.viii.1924, leg. Vinogra- dov; 1 ©, Kultuk, near Baikal Sea; 1 ©, Tunka, along river Irkut, about 180 km W-SW of Irkutsk, 20.vii— 10.vin.1911, leg. C. Ahnger, collection of Rodionov; 1 2, coast of Baikal Sea near Tolsty Mys, village Suk- hoy, vu.1928, leg. Vereshtshagin & Tikhomiroy; 1 9, Baikal, delta of the river Selenga, near the Proval bay, 12.vu.1925, leg. Vereshtshagin & Tikhomirov; 1 9, Kuznetsk basin (Kemerovo Prov.), river Suriekovaya, 2.vin.1951 (in coll. Leningrad and Wageningen). Additional material not seen by me. After reading my manuscript Dr P. Lindskog wrote me that he had examined an apparently undescribed species from a collection of Saldidae from Mongolia, which appears to fit the description of M. simulans. The specimens concerned were attributed to M. rivu- laria by Hoberlandt (1971b); they were collected in a small mountain range in NW Mongolia, SE Uvs- Nuur. The locality label reads as follows: Uvs Aimak, Somon Ondérchangaj, 1900 m, loc. 1090, 3 6 2 9, 11.v11.1968, Exp. Dr Z. Kaszab, 1968. Dr I. Kerzhner tested my key and could trace additional material of M. simulans in the Leningrad Museum. These data, which are also plotted on map 2, are as follows: W. Siberia: Bayan-Ölgiy aimak, river Ikh-Dzhargalan- tyn-Gol, 20 km NW of Bulgan, 1 d 2 9, 23— 24.v11.1978, leg. Gurjeva; same aimak, river shen yz-Agatsh-Gol, 15 km SE of Delun, 21.vii.1978, 6 ó 8 2, leg. Gurjeva; Bayan-Khongor aimak, river Tuin- Gol, 10 km S of Erdene-Tsogt, 7 d 16 ©, 25.vili.1978, leg. Gurjeva. Mongolia: Dzabkhan aimak, Dzegistay pass, 1 d, 23.1x.1926, leg. Kiritshenko (as M. rivularia in Vinokurov 1979b); Ara-Khangai aimak, confluence of Sumiyn-Gol and Tshulutyn-Gol, 5 d 72 (together with M. rivularia), 29—30.vi.1975, leg. Gurjeva & M. Kozlov. Comparative notes. — The distribution of this new taxon (see map 2) supports the view that M. simulans warrants the status of species, — distinct from M. rivularia. The latter species has been recorded from Siberia (even from the ex- treme north near the delta of Enisey), Mongolia (Hoberlandt, 1971), and Alaska. I saw addition- al material of M. rivularia in the Leningrad col- lections from the following localities: Mountain Pektusan, North Korea, 10 spec., 21.viii.1950, leg. Borkhsenius; Omsuktshan, Kolyma, 1 ©, 27.vili.1953, leg. Kurnakov; river Ebeten, 10 km SW from Kuysyur near Lena, Yakutia, 2 9, 10.vii.1957, leg. Gorodkov. Mongolia, Urga (Ulan-Bator), 7 spec., 4.vii.1926, leg. Kiritshen- ko; idem, 11 spec., 25.vi.1928, leg. A. Ivanov; Urga, coast of river Tola, a long series, 25.vi.1928, leg. A. Ivanov; Over Changai-ai- mak, Orchon waterfall, 2000 m, 1 dé, 14.v11.1965, leg. Muche (labelled as oorogallien Kir.). The presently known distribution of M. rivularia is shown on map 2. Future collections may perhaps reveal that the ranges of both spe- cies overlap partly. Within the genus Macrosal- dula, M. simulans, M. rivularia and the Alaskan M. monae form a separate unit, characterized by a similar median endosomal penis sclerite (fig. 11c); this sclerite is more slender in all oth- er Macrosaldula species (fig. 10g). Macrosaldula violacea sp. n. (fig. 11 i—k) Description. — For measurements, see table 1. Rather large (5.2—7.0 mm), slender, more or less parallel-sided, glossy black with metallic-vi- olet shine, wings inclusive of clavus and mem- brane entirely immaculate, antennae and legs predominantly dark brown, pubescence very R. H. Cossen: Eurasian Saldidae 239 short and sparse. Far East of the USSR, Japan. Alaska. This species is smaller (4.5—6.4 mm), This unicolorous black species is very close to rather dull, and bears a thick, somewhat shaggy, M. koreana (Kiritshenko, 1912a). The differ- pubescence of semilong silvery hairs. The sec- ences are tabulated as follows. ond dull-blackish species, M. simulans, has just 240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 violacea koreana size in general somewhat smaller, particularly the material from Japan in general somewhat larger, 6.3—7.3 mm (n = 20) which ranges from 5.2—6.0 mm (n = 8) ratio pronotum 2.7—3.0 (n = 5) length/width ratio antennal 2.6 —2.7 (n = 5) length/head width upper side colour hair covering short, less dense colour é strongly shining, with blue-violet inner side of first antennal segment, 2.42.55 (n = 5) 3.0—3.1 (n = 5) less obviously shining, sometimes with faint bluish reflection slightly more dense (this character can only be evaluated by simultaneous comparison of both species always entirely black, as in © transverse swelling above anteclypeus, anteclypeus and maxillary plates yellowish © all these parts black, transverse always entirely black, as in é swelling and maxillary plate sometimes partly black paramere fig. 11g regularly curved (fig. 9a, b) Other less readily visible characters reveal a greater discrepancy between both species. Whereas the abdominal stigmata in M. koreana have a normal position on the sternites, those of M. violacea are located in the lateral connexival membrane (fig. 11k). The median endosomal sclerite of M. violacea (fig. 11j) resembles that of typical Macrosaldula species, while M. ko- reana fig. 9c) in this respect more resembles M. rivularia (fig. 11c). Holotype (6), Japan, Honshu, Izumi Tamagawa, Tokyo, 24.vi.1951, leg. H. Hasegawa (in coll. Wage- ningen). Paratypes: 8 d 4 ©, idem; Honshu, Oni- kobe, Miyagi, 1 & 12.viii.1977, leg. T. Nambu; Hok- kaido, Matsuneshiri, Nakatonbetsu, 12 ¢ 1 9, 26.viii.1977, leg. M. Tomokuni; USSR, 29 specimens, Vinogradorka, Primorskiy Kray, 9 and 14.vii.1929, leg. Kiritshenko; 1 9, river Sitsa (now Tigrovaya), Sutshan (now Partizansk) district, Primorskiy Kray, 18.vii.1926, leg. Rostovykh; 1 d, river Sudzukhe (now Kievka), 7.vii.1948, leg. Sharov. Paratypes in Leningrad coll., coll. Kyushu Univ., coll. Hasegawa, Linnavuori, Polhemus, and Wageningen. Comparative notes. — Both M. violacea and M. koreana cannot be confused with other members of this genus. The only known en- tirely black species of the Macrosaldula group 1s M. rivularia (Sahlb., 1878) from Siberia and been described above. So far the scarce data suggest (see map 1) that M. violacea may be dis- tributed on the mainland and islands around the Sea of Japan. The continental localities all lie north of Korea. Of M. koreana I have seen ad- ditional material from localities at roughly 1500 km NW of Korea (coast of the river Shilka, Sre- tensk, Transbaicalia, 3—19.vii.1928, leg. Kapus- tin; of the large species in the Leningrad coll. I have studied 23 d and 14 ©). Other localities of M. koreana (1 ®, River Koppi, 52 km W of Mouth, Sikhote-Alin Mts, 20.viii.1924, leg. Emelyanoc; 2 ©, Imperatorskaya Gavan’ (now Sovetskaya Gavan’), Khabarovsk region, 23 and 26.vii.1916, leg. N. Krylov) lie north of Korea and thus overlap the distribution of M. violacea. It is uncertain whether M. koreana occurs in Ja- pan. Farlier records from Honshu (Hasegawa, 1960) may probably refer to M. violacea. Calacanthia grandis sp. n. (figs. 15d, 17d) Description. — For measurements, see table 1. Large (single 2 measures 7.5 mm) with pro- portionally long antennae; dull, black with small testaceous wing spots, adpressed short lightish setae and some erect dark setae on head and pronotum; superficially resembling Chilox- R. H. COBBEN: Eurasian Saldidae N > (u UZ Fig. 13. a—g. Macrosaldula roborowskii. a, median endosomal sclerite, plane view (below), lateral view (above): ‚ paramere; c, left fore wing; d, penisfilum; e, right antenna; f, male grasping plate; g, pronotum. h, 1, M. varia- bilis, eunomy of left fore wing; h, variab. connectens; i, variab. variabilis. j, M. jakovleffi, pronotum. 242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 anthus species from the Himalaya or large-sized dark Macrosaldula species. China. Head: dull, black inclusive of mouth sclerites, basal half of labrum brownish; preocellar spot through narrow testaceous band along eye mar- gin connected with pale spot bearing frontal tri- chobothrial setum; setosity short, golden, ad- pressed, and with scattered erect dark setae as long as the six trichobothrial-like setae. Thorax: Pronotum 2.6 times wider than long medially, side margins almost straight; frontal edges pro- truded laterad, much wider than pronotal col- lar; callus rather flat, not reaching side margins, central pit transverse, with smaller pit left and right, posterior lobe of pronotum medially half the lenght of callus; rather densely covered with short adpressed golden setae and with few erect dark setae; black, pronotal lateral margin very narrowly testaceous; acetabula 1 and 2 entirely lightish, 3 only apically. Wing of single specimen macropterous (fig. 15d); exocorium subbasally foliaceous, some- what wider than base of endocorium, hypocos- tal lamina well-developed, without secondary ridge; margin of wing slightly infolded at the small polished area which is adapted to receive the coupling plate of the male; corium dull, black with seven testaceous spots (fig. 15d), cla- vus entirely black; membrane ochreous, veins brown, cells with brownish markings; short sil- very pubescence of corium moderately dense, somewhat shaggy. Extremities: Antennae long, ratio of segments: 1 : 2.41 : 1.40 : 1.57; segment 2 cylin- drical (2), thickness 0.4 times diameter of seg- ment 1; all segments dark-brownish, with short adpressed silvery setae, segment 1 in addition with semilong dark stiff setae along median side, segment 2 with some scattered semilong setae which are slightly longer than diameter of seg- ment, erect setae on segments 3 and 4 approxi- mately as long as the diameter of the segment. Legs: flat upper and under sides of femora largely dark-brownish, fore and back testa- ceous, with short silvery pubescence and scat- tered obliquely erect dark setae, also on front femur; base and apex of tibiae dark-brown, otherwise testaceous, tibia 1 with one dark ring in middle, tibiae 2 and 3 with series of dark patches, length of dark spines subequal to diameter of tibia; second tarsal segment yellow- ish, third segment brownish, ventral side of sec- ond tarsal segment of third leg with two rows of six spines. Genital structures of 9, spermatheca piriform, with proximal flange (fig. 17d), teeth of ovipositor blade sharply pointed (fig. 17f), apex of second gonapophysis tapering (fig. 17f above.) Material. — Holotype (?), China Balang, Wassu- land, W. Szechwan, Sankiangkow, 7.viii.1934, leg. Friedrich (in Leningrad Museum). Comparative notes on Calacanthia. The genus Calacanthia is characterized by the flattened first and second antennal segment in the male (fig. 16d, e). Only three species have been assigned to this genus, viz., C. trybomi (Sahlb., 1878) and C. alpicola (Sahlb., 1880) from the northern arctic regions (habitat de- scriptions in Lindskog, 1975), and C. tibetana Drake, 1954, from Tibet. My re-examination of the species originally described as Acanthia an- gulosa Kiritshenko, 1912, and listed under Telo- leuca by Reuter (1912) as well as Drake & Ho- — berlandt (1951) in their catalogue, has shown that it represents a true Calacanthia (a reference to this unpublished conclusion was made by Lindskog, 1975). The new species C. grandis from China shares with C. tibetana and C. an- gulosa the protruded anterolateral edges of the pronotum (fig. 16g). This character and the sharply notched ovipositor blades (fig. 17f) sep- arate this Himalayan group of species from the arctic pair of species (compare with figs. 16f and 17e). C. grandis sp. n may be on the average sig- nificantly larger than the other species; the sin- gle specimen at hand measures 7.5 mm. The largest size of the type series of C. angulosa is 6.0 mm (® submacropt.), whereas the type se- ries of C. tibetana, including macropters, varies from 4.25—5 mm. C. grandis has proportional- ly much longer antennae; the ratio length anten- nal segments 2—4/width pronotal collar is 3.84 (2.3—2.9 in C. angulosa, n = 5; 2.3—2.5 in C. tibetana, n = 3). The second antennal segment is also more slender than in the other two spe- cies; its apex is about 0.7 the widest diameter of segment 1 (subequal in the other species). It re- mains, therefore, questionable whether the typ- ical antennal character of Calacanthia, which is demonstrated so clearly in the male sex, applies also for C. grandis, males of which are not yet known. Other differences between C. grandis on the one hand, and C. tibetana and C. an- gulosa on the other are: dark pattern of extremi- ties and forewings predominating in C. grandis and less in the other species (fig. 15d and fig. 15 e—i, respectively); erect dark setae scattered all over the head in the former, only two erect setae R. H. COBBEN: Eurasian Saldidae 243 Fig. 14. Pigment variation of wings of Macrosaldula species. a, M. scotica; b, M. madonica; c, M. tadzhika; d, M. jakovleffi; e, M. nivalis; f, M. miyamotoi. g, h, lateral and frontal side of male head; g, M. nivalis; h, M. jakovleffi. 244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 on vertex in the latter two species. C. tibetana and C. angulosa, both from Tibet, reveal a close phenetic relationship, and I considered the pos- sibility of conspecificity. However, some dis- crepancies may speak well for a specific status of both, awaiting more material from other lo- calities. C. tibetana has a more sharply defined ‘wing pattern (fig. 15i), indeed resembling Telo- leuca species (e.g. fig. 15b) with the exception of T. kusnezowi. The spots on the wings of C. angulosa are less prominent and ash-black areas on corium and clavus prevail (fig. 15e—h). In the latter species, the basis of the exocorium is clearly wider than the adjacent part of the endo- corium, whereas these parts in C. tibetana are at most subequal. The reduction of the forewing in not fully winged specimens is more marked in C. angulosa (weakly semibrachypterous, in ter- minology by Cobben, 1960) than in C. tibetana (submacropterous). A pronounced difference in reduction of flight ability can be deduced from the development of the hind wing. Its length is subequal to the fore wings in C. tibetana, but in C. angulosa the hindwings are only stubs, not surpassing the level of the scutellum apex. Such variations in wing development are apparently not related to differences in altitude. The type series of C. angulosa was collected at an eleva- tion of 4300-4700 metres (I saw an additional ® from E Tibet, 4000 m, Shopando, Kham, 4.v.1936). The type material of C. tibetana ori- ginates from 5000 m altitude. Two other charac- ters may be of value to separate both species, premised that additional material will confirm constancy. In C. tibetana, and not in C. angulo- sa and C. grandis, I found the ® ventral wing margin posteriorly of the hypocostal lamina provided with weak transverse ridges (fig. 15c). The spermatheca of C. angulosa is more than three times as long as wide at its base (fig. 17b), whereas in C. tibetana it is spherical (fig. 17c). Although there is considerable intraspecific variation in the shape of the spermathecal bulb in Saldula species (Karnecka, 1974), the noted differences in Calacanthia being exactly the same in two specimens of each species checked, might prove to be of significance. The same counts possibly for the parameres (fig. 16b, C. tibetana; fig. 16c, C. angulosa) which could be studied in only one individual of each species. Teloleuca kusnezowi Lindberg, 1934 Additional data. — The eunomic variation of the wing pattern of 7. kusnezowi Lindberg, 1934, and other details of this species are pre- sented in fig. 20, since after the original descrip- tion based on one female from Khabarovsk no further records were published. Abundant new material reveals that the species is distributed in USSR as is shown on map 4, but also occurs in Japan. The localities in Japan and USSR, from which I have seen material, are as follows. Japan (Hokkaido) Berabonai, near Ashoro, 2 d 1 2, 8.vi1.1958, leg. S. Miyamoto (together with S. no- bilis); Aizankei, 5 d 5 2, 1 larva, 9.viii.1967, leg. A. Nakanishi; Tennin-kyo, 1 d 1 2, 1 larva, 27.vi.1967, leg. T. Saigusa; Yukomanbetsu near Mt. Daisetsu, 5 ó 5 2, 2.vu.1970, leg. H. Hasegawa. (Honshu) Mt. Ya- tugadake, 4 d 3 2, 18.v11.1939, leg. H. Hasegawa; Masutomi Kai, 3 6, 26—29.vu.1957, leg. S. Miyamo- to. The Leningrad Museum contains series from the following localities: Amur region: Korsakovo, river Amur W of Svobodnyy, 3 d 3 2, 24.vii.1959, leg. Kerzhner; Birsherta, river Zeya, 50 km of Blagovestshensk, 1 ?, 19—26.vi.1914,leg. Popov. Khabarovskiy Kray: Imperatorskaya Gavan’ (now Sovetskaya Gavan’), 1 ®, 26.vu.1916, leg. Krylov, from Kiritshenko’s collection; Ozerpakh, delta of riv- em noie, NOMME Mulshernayın! Primorskiy Kray: confluence of rivers Iman (now Bol’shaya Ussurka) and Tatyube (now Dal’nyaya), 1 2, 24.vi1.1913 (Buyanova); Vinogradovka, 27 d 10 9, 5—6.vin.1929, leg. Kiritshenko; river Suputinka (now Komarovka), 1 2, 26. vii.1935, leg. Samoylov; ibi- dem, 2.iv, 8.vii.1937 and 1 6, 22.vii.1937, leg. Rich- ter; ibidem, 1 d 3 9, 12—22.v11.1940, leg. Ivanov; ibi- dem, stony bed of the stream Egerskiy Klyutsh, 1 ©, 30.vu.1953, leg. Kurentsov; Vladivostok, 1 à, 27.1x.1932, leg. Rysakov; ibidem, Shamora (now La- zurnaya) bay, 12 ©, leg. Stepanow & Shutova; river Sudzukhe (now Kievka), 3 d, 12—15.vin.1948, leg. Sharov; river Peyshula (now Suvorovka), 1 9, 12.vi1.1963, leg. Nartshuk; Frolovka, Sutshan (now Partizansk) district 1 34 31.1926 and 272% 7.v11.1926, leg. Rostovykh; Tigrovaya, same district, 3 d 5 9, 26.vi—3.v11.1928, leg. Rostovykh; Fanza (now Rutsh’i), same district 1 d, 15.v11.1926, leg. Rosto- vykh; Derzhanovo, same district, 1 6 1 Q, 12.v11.1928, leg. Rostovykh; Sedanka, Vladivostok, 1 9, 20.v1.1927, leg. Sokolov. Salda kiritshenkoi sp. n. (figs. 18a left, b, c, 21d) Description. — For measurements, see table 1. Large (5.2—6.8 mm semibrachypterous, up to 7.8 mm macropterous), rather dull, coal- black species with fine cover of very short, densely packed brown setae. East-USSR, Japan, NE China. Since the resemblance with Salda mueller (Gmel.) is very strong, only the differences will be mentioned. Male genital structures (fig. 15b, c) are of the R. H. COBBEN: Eurasian Saldidae 245 Fig. 15. Left forewing. a, Macrosaldula nivalis; b, Teloleuca brancziki; c—j, Calacanthia species; c, 1, Calacan- thia tibetana, c, underside of semibrachypterous specimen, with hyposcostal lamina and coupling pit (enlarged in left figure), i, macropterous specimen; d, C. grandis, holotype © ; e—h, C. angulosa (cross-hatching on cori- um and clavus indicates ashy-black colour); j, C. trybomi. 246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 kiritshenkoi ratio length/width 1.7—2.0 (semibrachypterous) lateral wing margin short brown hairs on upper side of thorax and wing strongly convex (fig. 15a left) dense (fig. 18d), giving the species a more dull appearance muelleri 2.0 22 less arched (fig. 15a right) very sparse (fig 18b) corium without punctations punctated corial veins hardly visible distinct colour maxillary plates black, anteclypeus mentioned parts in general more with narrow, light basis, only extensively ochreous lightish midpart of labrum lightish, first antennal segment and tibiae tending to blackish head with only few short hairs somewhat more hirsute same shape in both species. The most distinctive constant difference is the layer of short setae on the wing, which is much more dense and very regularly spaced and orientated in the new spe- cies (fig. 18d). I could not find any morphome- tric difference, except for the on the average broader shape of short-winged S. kiritshenkoi. All specimens lack any light spot on the corium, whereas in S. muelleri the mesocorium sometimes has one white spot. S. morio Zett., also a related species, has usually several endo- corial spots, and is otherwise recognizable by the more or less shiny, not punctated cuticle. The only North American Salda species with which S. kiritshenkoi could be confused is S. buenoi (Mc Dunn). However, S. buenoi has part of the lateral wing margin almost straight, a more dense and longer golden pubescence, cori- um with light spots and longer distal processes of the median endosomal sclerite; such a pro- longed sclerite is an apomorphic character of all other N. American Salda species, except for the Holarctic S. littoralis (L.). Length of macropterous holotype ¢ 6.0 mm, width 2.8 mm. Length of paratypes varying from 5.2 mm (semibrachypt. d) to 7.8 mm (ma- cropt. 2). Material. — Holotype (d) (macropterous), USSR, Primorskiy Kray: valley of the river Odarka, about 25 km from Station Evgenevka (city Spassk-Dal’niy), 16.vi (old style calendar, = 29.vi), 1911, leg. A. Tsherskiy (in Zool. Mus., Leningrad). Paratypes, USSR, Primorskiy Kray, 1 2 semibrach., 2 2 ma- cropt., idem 4. (= 17).vii.1911; Sivakovka, south shore of lake Khanka, 2 9 semibrach., 23.vi.1924, leg. Samoylova; lake Khanka, Kamen’-Rybolov, 1 4, 30.vi.1910, leg. Tarobarov; Tshernigovka, 1 © semi- brach., 5.vii.(= 18.vii).1912, leg. Emelyanov; Ryaza- novka, Khasan district, 3 d 1 © semibrach., 9.vii.1982, leg. I. Kerzhner; Vladivostok, Station Okeanskaya, 1 2 macropt., 10 (= 23).vii.1911, leg. Stshavinskaya, from the coll. of Kiritshenko. Khabarovskiy Kray: Krasnaja retshka, pr. Khaba- rovsk, 3 9 semibrach., 2 © macropt., 2 d semibrach., 10.vii.1931, leg. Pereleshina; Knyaze Volkonskoye, nr. Khabarovsk, 8 d semibrach., 4 & macr., 5 2 semi- brach., 7.vu.1977, leg. Stys & Davidova (dry salt- steppe); Khabarovsk-Ussuri, 1 d& macropt. 1.vii.1978, leg Styps & Vilimova; Khabarovsk south, 1 2, 4.vu.1977, leg. Sys & Davidova. Kunashir Island: Golovnin volcano, 2 © semibrach., 10.vii.1980, leg. A. Gorokhov; Sernovodsk, Glukhoe lake, 1 6 1 2 semibrach., 28.viii.1980, leg. Egorov & Kanyukova. NE China: railway stations Shanshi and Hailing be- tween Harbin and Mudankiang, 1 © semibrach., 7.vu.1902, leg. Krylov. Japan: Akkeshi, Hokkaido, 4 9 semibrach., 5. vii. 1958, leg. T. Nakane. Paratypes in Zool. Mus Leningrad, Jap. Nat. Museum and in coll. Wageningen, Hasegawa (Japan), Lattin (Ore- gon), Polhemus (Colorado), Popov (Moscow) and Stys (Czechoslovakia). I have further seen a specimen (1 2 semi- brach.) labelled: Proskurov (Khmel’nitskiy), vii.1895, leg. Zubovskiy; this locality is in the Ukraine, far away from the territories near the Sea of Japan. Future collections may prove whether the Ukraine indeed lies within the range of the species, or that the specimen in question has been mislabelled. A recent collec- tion received from Dr Stys confirms that S. kiritshenkoi occurs much further to the West than expected. The new material is from Kirgi- R. H. COBBEN: Eurasian Saldidae zia, Ala-Archa, 2000 m, near Frunze, 2 d 2 9, 21.vi.1982, leg. Stys (very humid habitat along stream, on open places in between thick vegeta- tion of moss and sedges). The only difference with the Far-East specimens is the fact that the central-Asian material has somewhat more pro- nounced elytral veins, lighter coloured hind tib- iae and graphite-black membrane. For comparative notes, see next chapter. DISCUSSION OF EURASIAN SALDA SPECIES The records of S. kiritshenkoi (map 3) suggest a broad-band distribution in Asia between 43° and 61° latitude. The labels do not always in- clude data on altitudes, but the occurrence around the lake Khanka and the mentioned lo- cality on Hokkaido suggest it to be a lowland species, although the Kirgizian population was found at 2000 m altitude. The closest relative, S. muelleri, also lives predominantly at low and submontanous altitudes. The distribution-map of this species so far shows only few scattered localities in the USSR (map 3), but it has a very vast, probably continuous distribution from West-Europe on to the Sea of Japan. I saw 1 © 247 from Khabarovsk collected together with a se- ries of S. kiritshenkoi (leg. Stys & Davidova, 7.vii.1977). Other species which could be confused with both S. kiritshenkoi and S. muelleri at superficial inspection are S. morio (Zett.), S. micans Jak. and S. splendens (Jak.). Since I could study the type material of the latter two poorly known species, further data on them are presented here. The main differences of semibrachypterous specimens can be tabulated as follows. When S. kiritshenkoi is added to the left of this table, we have a series of species with de- creasing pubescence and increasing polished cuticle from left to right. S. morio, micans and splendens show a weak metallic lustre of the dorsal surface. S. splendens is easily recognised by erect semilong dark setae on thorax and hemielytra (fig. 16d). Identification of single in- dividuals of S. muelleri, morio and micans may be difficult and the combination of tabulated characters may help the decision. There is a great overlap in body size and width but the mean value (underlined) of S. muelleri (20 3, length in mm 4.73—5.21—5.80, width 2.21— muelleri morio micans splendens hemielytra: texture punctate, weakly impunctate, more or polished, strongly polished, strongly shining less distinctly shiny, shining shining except for exocorium and membrane hemielytra: setosity with sparse, short very scattered short as S. morio erect semilong dark setae (fig. 18b) setae (fig. 18a) setae ın addition to sparse short setae (fig. 16d) hemielytra: corıum rarely with corium often with entirely black entirely black pigmentation one midapical light some light spots; spot; membrane of inner margin of macropter for major membrane usually part dark (fig. 16k) lightish, membrane of macropter for major part lightish (fig. 16j) first coxa black, apex black, apex often entirely pale entirely pale sometimes lightish lightish pigmentation of fig. 16a fig. 16b fig. 16c fig. 16c femora pigmentation of first fig. 16e fig. 16f fig. 16g fig. 16g and second antennal segment RS MU SLA VR NE RP D Te ENE ET EE IRE U SINO RATA RA 248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Fig. 16. Calacanthia species. a, f, h, C. trybomi; b, C. tibetana; c, €; 8» C. angulosa; d, C. alpicola. a, subgenital plate 2; b, c, left paramere, b, innerside, seen in direction of arrow in b; d, e, right antennae; f, g, pronotum; h, head, Goal aspect. R. H. COBBEN: Eurasian Saldidae 249 2.39—2.79; 17 ©, length 5.30—5.98—6.55, width 2.73—2.98—3.20) is smaller than the re- spective dimensions of S. morio (18 d 5.0— 5.84— 6.30, 2.53—2.63—2.80; 20 9, 6.08— 6.54—6.96, 3.00—3.19—3.55). The material of S. micans and S. splendens is too limited to allow reliable comparison, but these species tend to fall into the size class of S. muelleri (S. micans, 3 3 length 4.5—4.9, width 1.8—2.1, 2 2 length 5.2—6.1, width 2.36-3.0. S. splendens, 3 3 length 4.4—4.8, width, 1.7—2.0, 2 ® length 5.7—6.5, width 2.6—3.1). I could not find reliable differences, neither in the genitalia (the median penis sclerite seems to be more slender in S. micans and splendens (fig. 15j), but this should be checked in more speci- mens), nor in the many ratios calculated from a variety of measurements. Considerable varia- tion of ratios exist, which is partly due to allo- metric differences between smaller and larger individuals. The general facies of some smaller specimens of S. splendens and S. micans looks at first sight somewhat different from the other species (pro- notum with straight lateral margins, humeral edges more acute and somewhat upturned). Other specimens are more like S. muelleri and S. morio, in which the shape of the pronotum also varies. The metallic lustre mentioned in the original description of S. micans is not consis- tently present in the series seen by me, but often also occurs in S. muelleri (according to Wroblewski, 1966) and in S. morio. Whereas S. muelleri and S. morio from West Europe can be reliably discriminated, specimens from the eastern Palaearctic are more difficult to identify, especially the long-winged forms in which the dorsal cuticular texture tends to be intermediate. In macropters of S. morio from the USSR (Irkutsk, leg. Jakovlev, 3 d 2 9, to- gether with semibrachypterous specimens) the wing is less shiny, and of S. muelleri (4 © from different localities in USSR) less punctate than in typical specimens. However, the difference in dorsal pubescence between both species remains constant; this is also mostly true for the differ- ence in pigmentation of the membrane (fig. 16), k). Since I had no opportunity to reidentify material of all published records from eastern regions, the distribution patterns of $. morio and S. muelleri remain somewhat uncertain (possible misidentifications). The identity of some specimens (see below) remains dubious until more material will be available for study. The available information is summarized below under the individual species. Salda muelleri (Gmelin) This species occurs further south in Great Britain (Scudder, 1958) and all over the conti- nent than the next species (S. morio). In the me- diterranian subregion it has been recorded inci- dentally from France (Puton, 1880). Greece (Reuter, 1895) and Turkey (Lindberg 1922). In Poland it is known from some 20 localities (Wroblewski, 1966) and in Czechoslovakia it seems to occur in northern localities of the Car- pathian Basin (Benedek, 1970) and in Bohemia (Stys, unpublished). Both this and the next spe- cies have not yet been collected in Albania, Yu- goslavia, Bulgaria and Roumania (Josifov, 1970). To the east, the exact distribution pattern is poorly known. Its occurrence is broadly indi- cated by Kiritshenko (1951) as: “Forest zone of the European part of the USSR up to Volyn’, Khar’kov and Ryazan’ provinces”. Vinokurov (1979a) listed it for East Siberia, and Kerzhner (1978) as a questionable record for Kunashir Is- land which, however, appears to belong to S. Riritshenkoi. I have seen the following material, mostly from the Leningrad museum (semibrach. if not otherwise stated): Leningrad region: Ol’gino, 3 d 9 2, 6 larvae, 20.vi.1901, 17.vi—9.viii.1902, leg. Bianchi; Krupeli, 1 2, 30.vi.1897, leg. Mazarakiy; Terioki (now Zeleno- gorsk), 2 2, 1889, leg. Wagner; Shuvalovo, 1 6, 25.v.1897, leg. Zubovskiy; Chernaya Lakhta, 1 9, 1.vii.1904, leg. Bianchi; Log near Luga, 1 6, 1 larva, 18.vi—1.vu.1918, leg. Jacobson; Sablino, 1 d 1 ©, 21.v1.1921, 6.vi1.1922, leg. Bianchi; Ostrovki, river Neva near Schüsselburg (now Petrokrepost’), 2 d 1 @ macr., 5—6.vi. 1906 (leg. Jacobson); Lobanovo, 1 9, 14.v1.1906; Lakhta (now part of Leningrad), 1 ®, 22.v1.1919, leg. Reichardt; Pomeranye, 2 d 4 9, 10.vii.1911, leg. Ihgin; river Tigoda, 4 d 1 Q, 4.v1.1911, leg. Semenov - Tjan - Shanskij (series of this locality somewhat less punctate); Svir, 1 ©, (?), leg. Günther; Listy Nos (now part of Leningrad), 1 2 macr., 20.vi.1889, leg. Silantyev.; Zelenogradskaya railway Station, near Moscow, 1 2, (?), leg. Y. Zhe- zekhin (in coll. Popov). Karelian SSR: Petrosavodsk, 181%, (?), leg. Günther; Muromli, 1 d 1 © (?), leg. Günther; Vitebsk region: Vitebsk, 1 ©, (?), leg. Biru- la. Arkhangelsk region: Shipitsino, 3 d 3 2, 1 2 macr., 1 larva, 20.vi and 6.vii.1942, leg. Stark (note of Dr Kerzhner: “at my experience this label is wrong and refers to insects collected in more southern re- gions”). Pskov region: Kharlamova gora near Gdov, 1 3 1 ©, 15.vi.1898 (leg. Bichner). Kalinin region: Bo- logoe nar Sey ar va 5 vin 1908 10.71.1904; 11.vin.1904. Estonian SSR: Gapsal (now Haapsalu), 1 250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 2, (2), leg. Morawitz; Sillamyagi (now Sillamyae), 1 2, 25.vi.1930, leg. Bianchi; Merrikul, 1 d, 7— 10.vii.1904, leg. Somira. Volgograd region: Sarepta (now part of Volgograd), 2 6, (?), leg. Becker, Rya- zan region: Kazachiy near Rannenburg, 1 9, 24.vi.1903, leg. Semenov. Ukraine: Rovno region: Krasnoe, near Dubno, 1 & 1 ®, (?), leg. Karavaev; Chermyakovo near Ostrog, 1 ®, 1—5.vii.1900, leg. Neklyudov. Volyn’ region: Zamostochye near Senki, 1 2, 14.vi.1905, leg. Birula. Khmel’nitskiy region: Kamenets-Podolskiy, 1 ©, (?), leg. Birula. Donetsk region: Yarovaya near Svyatogorsk (now Sosnovo), 1 3, 19.vi.1938, leg. Arnoldi. Kazakhstan: Aktyubinsk region: Berchogur, Mugodzhazy Mts, 1 9, 8.vii.1932, leg. Lukjanovitsh. Turgay region: Kokshe- tau Mts, 1 ©, 23.vi.1957, leg. Asanova. Siberia: Ir- kutsk region: river Belaya, tributary of Angara, 2 d 1 ©, leg. Gartung. Maritime Province Far East, Primorskiy Province, Knyaze Volkonskoye near Khabarovsk, 1 ® macr. (together with S. kiritshen- kot), 7.vii.1977, leg. Stys & Davidova. Salda morio (Zetterstedt, 1840) As far as it can be concluded from the avail- able reliable records this species has a more dis- tinct northern Eurosibirian distribution than the foregoing species. Its range extends eastward into N. Mongolia (Josifov & Kerzhner, 1967, Hoberlandt, 1971a, Vinokurov, 1979b), Kuril Isl. (Kerzhner, 1978) and Japan. I have verified Russian materal from the following localities: Karelia, Muromli, 2 ©, leg. Günther; idem, Ladoga, 1 ©, leg. J. Sahlberg; Ukraine, Krasnoe, Volyn’ region (now in Rovno province), 1 2, leg. V. Karavaev (to- gether with S. muelleri); environment of Irkutsk, Pashkovskoe, river Angara, 1 6, and Markovo, 2 d 1 2, leg. Jakovlev; Mongolia East aimak, river Nömrögin-Gol, 32 km SE of Mt. Salkhit, 1 d 1 9, 8.vi.1976, leg. I. Kerzhner; East aimak, Mt. Derkin- Tsagan-Obo, 60 km ENE of Bayan-Burd, 1 d 19, 3.viu.1976, leg. I. Kerzhner; Transbaicalia, river Ingo- da, 1 2, 11.vii.1989, leg. G. L. Suvorov (type locality of S. splendens); Amur region, Samodon near Korsa- kovo, 1 ©, 7.viii.1959, leg. I. Kerzhner; Klimoutsy 40 km W of Svobodny, 1 & 1 2, 15.vii.1959, leg. I. Kerzhner; Primorskiy Kray, Ryazanovka, Khasan district, 3 d 1 9, 9.vii.1982, leg. I. Kerzhner; Sagha- lien, Shiritori (now Makarov), 1 d 2 9, 3.vili.1938, leg. H. Hasegawa. The only published record of S. morio in Ja- pan is the Oze district, Honshu (Asahina & Ha- segawa, 1951, and Hasegawa, 1954). I have seen long series from this region and can conclude only provisionally that this belongs to S. morio, despite of some minor differences with Euro- pean specimens (corium between veins less shining). I further studied 2 & and 1 @ from Hokkaido, Mt. Daisetsu, 8.v111.1967, leg. A. Nakanishi. Dr Lindskog (Stockholm) kindly sent me for judgment 1 & and 1 ® collected by Poppius near Ytyk-haja, Lena River (in Zool. Mus., Hel- sinki), identified as S. morio by H. Lindberg. As Lindskog already noticed both specimens have a very polished dorsal surface, the outer margin of the exocorium being dull. The highly shining wings, without erect setae, are reminiscent of S. micans, but in contrast to this species, the cori- um bears lightish spots and the membrane is largely unpigmented. Both rather large speci- mens (d 5.9 mm, © 6.5 mm) are somewhat ten- eral. Provisionally, I must conclude that they belong to S. morio, awaiting larger samples from that region. Salda micans Jakovlev, 1889 Five specimens are located in the Leningrad Museum, one of which (9) from the type locali- ty: Kultuk (leg. B. Jakovlev). I designated this specimen as the lectotype with an appropriate red label. The data of the other specimens seen by me are as following: Tuva (Tuvinian autonomous province): Shagonar forestry, river Ulug-khem (= Yenissey), locality Ad- ar-khysh, 2 d 1 2, 2.vu.1956 (leg. Levin); Yakutian (Yakutian ASSR), Badarannakh, ca. 100 km W of Ya- kutsk, 1 d, 17.viii.1926, leg. Ivanov. These localities are plotted on map 3. Salda splendens (Jakovlev, 1905) Five specimens from the type locality are lo- cated in the Leningrad Museum (Transbaikal, river lnsodar 2 OG 3 So Hy 898 les. G. Il. Suvorov; 1 ® has no locality label, but the same collector label). I designated one of the males as the lectotype with an appropriate red label. The following new additions can be mentioned: Tunskinskie Gol’tsy Mts, E. Sayan mountain region in Buryatian ASSR, E. Siberia, 1 2, no other data; erect hairs somewhat rubbed off (in Coll Mus. Nat. Hung.). Badarannakh, ca. 100 km W of Yakutsk, 16.vin.1926, leg. Ivanov (pin with only rostrum and one forewing); this locality is the same as one given for S. micans, but collected one day earlier; N. Korea, Daitaku, Kankyo-hokudò, 2 6, 8.vii1.1939, leg. M. Tanaka (1 d in Ent. Lab. Kyushu Univ. Japan, 1 d in coll. Wageningen). Localities are plotted on map 3. Salda littoralis (Linnaeus, 1758) Also on the status of subspecies pzechocku Wagner, 1967, and of S. nevadensis Wagner, 1960. R. H. COBBEN: Eurasian Saldidae 251 Dr OS Fig. 17. Calacanthia species. a, e, i, C. trybomi; pina (Ce angulosa; c, f, g, j, k, C. tibetana; d, f, C. grandis; h, I, C. alpicola. a—d, spermatheca; e, f, right view of ovipositor gonapophyses; g—i, parandria; j, right grasping plate of d ; k, 1, median endosomal sclerite. 252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 S. littoralis is the most wide-spread represen- tative of the genus in the whole of Europe and Palaearctic Asia. The material which I have seen from Japan (first recorded for this country in 1968, Hiura), Alaska, Canada, N USA (16 spec- imens from eight localities), indeed conforms entirely in external and internal structures with specimens from Europe. From the abundant material present in the Leningrad Museum and Popov’s collection (Moscow), partly plotted on map 3, we may conclude that the species occurs everywhere in the USSR where suitable habitas occur (marine and inland salt-marshes, exposed fresh-water swamps in mountainous areas). The next to no data from W. Siberia may be due to paucity of collections made in that region (Kerzhner, correspondence). The remarkable ecological duality of S. littoralis is most promi- nent in W. Europe where the species is restrict- ed to salines along the Atlantic coast and to mountains from 1800 up to 2300 metres altitude (the Alps, Heiss, 1972; the Pyrenees, person. observ.). It is remarkable that S. hittoralis seems to be absent in the Carpathian mountains (Ben- edek 1970) loberlandra 19774 2Stys, pers: comm.). On the other hand, this mountain chain everywhere harbours the endemic saldid Teloleuca brancziki, which lives on flat finely graveled banks of larger streams. S. littoralis does not inhabit such localities, so that its ab- sence in the Carpathians can not be simply ex- plained through displacement by 7. brancziki. The most southern inland-population in W. Europe, the Sierra Nevada, Spain, was described as a valid species: Salda nevadensis Wagner, 1960. However, I could not confirm the stated differences between this species and S. littoralis. The latter species should have larger eyes, the vertex being only 0.85—0.90 (d) and 0.95— 1.02 (2) times as wide as one eye. The head and especially the gula should be longer, the labrum larger and more porrect, the pronotum and forewings wider than in S. nevadensis. My mea- surements, based on eight male and 14 female paratypes of S. nevadensis and on a double number of S. littoralis from various origins in the NW and NE Palaearctic, do not show diver- gence in the characters mentioned. For exam- ple: the ratio head width/width of vertex varies in S. littoralis between 2.41—2.54 (d), 2.22— 2.39 (2), and in S. nevadensis between 2.46— 2.50 (4), 2.33—2.39 (9). The ratio body length/width of S. nevadensis: 2.13—2.24 (6d), 1.97—2.15 (2), also falls within the range of variation in S. littoralis: 2.07—2.23 (6), 1.94— 2.15 (2). The only remaining actual difference between S. littoralis and the single population of S. nevadensis is the overall smaller body-size of the latter; reduction in body dimensions is not surprising since the type locality of S. nevaden- sis is at nearly 2900 m altitude. However, this discrepancy is not so pronounced as stated by Wagner. For his new species he mentioned a length of 4.7—5.1 mm for the mais and 4.9— 5.4 mm for the females. He contrasted these numbers with 6.0—6.5 (d) and 7.0—7.5 (@) in S. littoralis. However, according to the litera- ture (Cobben, 1960), the length of S. littoralis varies between 5.1 and 6.0 mm (6) and 5.9 — 7 mm (?) in the semibrachypterous form. The lengths of males of 6.0 up to 6.5 mentioned by me in 1960 applied to macropters, but since the nearly sixty original individuals of S. nevadensis are all semibrachypterous, comparisons should . be made with that morph only. Finally, Wagner (1960) stressed the fact that, whereas male and female of S. littoralis fall in clearly different size classes, the sexes of S. nevadensis can only be separated by checking the genitalia. Judging from the sample I have seen, this seems some- what overstated. The 14 females at hand vary from 5.4—5.7 mm, and the males from 4.8—5.0 mm. After all these refutations I must conclude to the synonymy of S. nevadensis with S. litto- ralis (syn. nov.). It is submitted that this is done purely on morphological grounds, but we may not expect that proof or disproof of reproduc- tive isolation between the Sierra Nevada popu- lation and other populations will be supplied in the near future. The description of S. littoralis piechocku Wagner, 1967, from Mongolia (1160—1750 m) suffers likewise from discrepancies with the real situation. This subspecies of about equal size as the nominate form, should have a more slender body shape, and shorter antennae and legs. The type material, which I could study (5 4, 3 ©), conforms in these respects with typical S. litto- ralis. The subspecies piechocki should have four windings of the base of the penisfilum, and the nominate subspecies 5. I checked the 5 males on this character, but I found the basal length of the filum fit within the variability range of S. lit- toralis. In my view there is no valid criterium to uphold the subspecies taxon piechockit. The coastal subpopulation of S. littoralis seems to have undergone at least one speciation process somewhere along the Adriatic Sea. De- viating material was originally described as a va- riety of S. littoralis, viz. adriatica (type locality R. H. COBBEN: Eurasian Saldidae 253 Fig. 18. Salda species. a, anterior part of corium of left fore wing, outline of lateral margin and vestiture of small setae; left, S. kiritsjenkoi; right, S. muelleri; b, c, S. kiritsjenkoi; b, median endosomal sclerite; below plane view; above, left lateral view; c, paramere; d, e, S. morio; d, median endosomal sclerite; e, paramere; f—h, S. micans; f, parandria; g, paramere; h, penisfilum; i—n, S. splendens; i, male grasping plate; j, median endosomal sclerite; left, plane view; right, left lateral view; k, base of penis filum, length of three individuals; I, apex of paramere; m, parandria; n, parameres; left, specimen from N. Corea; middle and right from type locality. 254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 “Illyria”) by Horvath, 1887, and subsequently (1901) recognized by himself as a valid species. Specimens studied by me at the time from the Gulf of Venezia indeed are characteristic enough to warrant specific status (Filippi, 1957; Cobben, 1960). S. adriatica has subsequently been reported from Greece and Bulgaria (Josi- fov, 1961, 1970). The population structure of this group of shore bugs in the Balkans and Asia Minor became, however, more complex after the description of S. subcoriacea Horvath, 1901 (type locality, Turkey, Aydinh, close to the Marmara Sea). To judge from the description, this Salda is more or less intermediate between typical littoralis and adriatica, and therefore I treated it, in 1960, provisionally as a subspecies of S. littoralis. As such were also identified spec- imens from Turkey (Hoberlandt, 1948) and Greece (Josifov, 1970). A deliberate decision on its taxonomic status and its relation to the S. /it- toralis-adriatica complex must be postponed until long series are sampled at regular intervals along the coastal lines of the Adriatic, Aegean, Black and Caspian Seas. The saisonality should also be considered. Subcoriacea-like adults were collected by me in a coastal swamp in Greece (near Khalkis, 29.vii.1978). I brought them liv- ing to Wageningen, hoping that these animals from 30.5° latitude would be easier to rear than S. littoralis from higher latitudes and altitudes. This species is univoltine in W and N Europe and has an obligatory winter-diapause in the egg stage lasting nine months (Jordan & Wendt, 1938; Cobben, 1968). This makes it very diffi- cult to rear large numbers in succession. My purpose was to try to hybridize the Greek pop- ulation with S. littoralis from the Netherlands and the Pyrenees. However, it appeared that al- so the animals from Greece went directly into a strong egg diapause under normal laboratory conditions. Such a phenology is reminiscent of a northern origin of the ancestor of S. littoralis. It would be interesting to know whether the true S. adriatica of the Venezia lagune still has re- tained this rigid life cycle. TENTATIVE KEY TO MACROSALDULA SPECIES Some historical remarks on the ranking of the taxon Macrosaldula as a genus were given on page 232. Most species ot this group are exter- nally recognizable by heteropterists familiar : with shore bugs by their rather stout size, slen- der and mostly dark coloured habitus with pro- portionally long antennae without long erect se- tae on the second segment, and by their lapidi- colous and agile behaviour, preferably along streams. The combination of these characters separate them from the smaller sized species of typical Saldula, a cosmopolitan genus most rich in species, and from some less typical Saldula subgroups, among which the orthochila group (page 226). The limits of both supraspecific taxa Macrosaldula and Saldula will be phylogeneti- cally redefined elsewhere; in such a study other less obvious characters will also play a role. Some Macrosaldula species strikingly deviate in colour (inornata, heyningent), or in colour and swollen antennae (roborowsku, see below) from the other group members. Three species deviate in some less readily visible, but important char- acters, such as: shape of hypocostal wing mar- gin, position of stigmata, and shape of median endosomal sclerite. These species are koreana, monae and rivularia, of which the generic posi- _ tion may need adjustment later. It is only for practical reasons that I include all 21 species listed in the following key as belonging to Ma- crosaldula. The key includes also one species which up till now has not been associated with Macrosaldula-like species, viz., the species de- scribed by Jakovlev (1890) as Salda roborowsku from the western part of China (“Chinesian Turkestan”). It was placed in Chartoscirta by Reuter (1895), Oshanin (1912), Hoffmann (1933), Wu (1935) and Drake & Hoberlandt (1951), spelled as roborowsk:). Dr Kerzhner in- formed me that the type series from “Oasises Nia and Keria” included more than one speci- men judging from the indicated length (4/— 4’ mm). At the present time the type speci- mens could not be traced in the Leningrad Mu- seum, but I could study another series located in that museum, apparently belonging to the same species. This material (1 & 2 2) was collected along the river Tisnaf, 6.vii.1890, leg. Grombtschewski. The locality is about 500 km W of the type locality and is plotted on the dis- tribution map 1. Both localities are, according to Kerzhner, along the old route passing be- tween the southern part of the Takla-Makan de- sert and adjacent mountains; the altitude is about 1500—2000 m. Examination of this material reveals that it represents a member of the Macrosaldula clade according to the elonga- tion of antennae and structure of the male grasping plate and genitalia. It lacks the two synapomorphies of Chartoscirta species (hind femur with stridulatory ridges, spherical median endosomal sclerite), but the swollen antennae (fig. 13e), somewhat elevated callus of the pro- R. H. COBBEN: Eurasian Saldidae 255 g Fig. 19. Salda species. a—c, left side of left femora 1—3; a, S. muelleri; b, S. morio; c, S. micans and S. splendens; d, S. splendens, left view of holotype; e—f, antennae; e, S. muelleri; f, S. morio; g, S. micans and S. splendens; h, i, paramere of S. micans; j, S. morio, Saghalien, Japan, pigment variation of wing; k, S. muelleri, macropterous wing. 256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 notum (fig. 13g), and the contrasting brown- white pattern of the corium (fig. 13c) are to some extent reminiscent of Chartoscirta, espe- cially C. dilutipennis (Reuter), occurring in Tur- kestan. I present in this paper also the illustrations of individual parameres of a number of Macrosal- dula species. The differences exhibited should not be regarded as absolute, since the intraspe- cific variability has to be tested on population level. In general, paramere morphology is of on- ly limited diagnostic value in saldids. The eu- nomic series of wing pattern shown in some il- lustrations must also be considered with some caution. I selected only those variations which reveal a smooth gradation from light to dark colour forms. Certainly, numerous small devia- tions from the ideal eunomy will be encoun- tered. For some species additional material may prove that the extremes of the light and the dark morphs will exceed the illustrations provided here. Distributions are given on a large scale. For detailed country records and data on ecolo- gy see Cobben (1960), Heiss (1972), Hober- landt (1977), Lindskog (1975), Vinikurov (1979b), and Wröblewski (1966). 1. Antennal segment 2 1/2 times and segment 4 about 2 times as thick as segment 2 in the middle (fig. 13e). Colouration of wing cas- taneous brown with two transverse white fascia (fig. 13c). Pronotum with pale mar- gin; callus somewhat swollen; (W China)... roborowskii (Jakovlev, 1890) comb. nov. — Distal segments of antennae not dilated. Colouration of wings otherwise ........ 2 2. Wings inclusive of clavus, and legs largely straw-yellowish. Lateral pronotal sides broadly pale (fig. 8); (Iraq) .. =) ackeprement prevallinca ss) 3 . Corium and clavus entirely devoid of light- ish spots. Wings with only adpressed short SETA PRE con EN RL. 4 — Corium and often clavus provided with lightish pattern. Wings with only adpressed short setae, or with erect semilong or long setae in addition to short pubescence .... 8 4. Dorsum shining. Inconspicuous pubescence regular, very short and adpressed. Vertex and pronotum without erect semilong setae DRG EA TERA EARLY, OEE NO STE 5 — Dorsum dull. Adpressed short pubescence more dense and conspicuous. Vertex and pronotum with some erect dark setae al- most as long as the six cephalic bristles ... 6 GQ inornata sp.n. 5. Strongly shining with blue-violet reflection (for further differences with the next species, see p. 240); (Far East of the USSR, Japan). 2 SEREEN violacea sp.n. — Less obviously polished, deep-black, sometimes with faint bluish reflection; (Far BastortheUSSR,Korea) ser zen I SION RE koreana (Kiritshenko, 1912) 6. Middle pair of cephalic trichobothrial setae arising from pale spot (for further differ- ence with next species, see p. 238) (Siberia, MongoliayAlas kala sea at ea een rivularia (Sahlberg, 1878) — Middle pair of cephalic trichobothrial setae atisinstomnblacktcuricle nnen 7 7. Ratio antennal segments 2 + 3 + 4/width of pronotal collar above 3.6. (USSR, region Krasnoyarsk-Irkutsk) ...... simulans sp.n. — Ratio antennal segments 2 + 3 + 4/width of pronotal collar below 3.6: dark extreme of | variabilis variabilis, and possibly other spe- cies of which the extreme melanistic form is notyet known. aac ur La eee 8 8. Corium with large subapical orange mark on exocorium; otherwise unicolorous black; (Italy Spain) ee eee si UES ae. heijningeni (Cobben, 1959) — Corium with varying lightish pattern .... 9 9. Pronotum with pale lateral margins. Middle pair of trichobothrial-like cephalic setae arising from pale spot (fig. 7d) ......... 10 — Pronotum and frons of head entirely black ee Se ET ARES ER GS Sco ola 11 10. Clavus proximally with longitudinal light- ish spot (fig. 7a); dorsal pubescence not denseyeoldent|(lranscaucasia) nee SEU Lin un clavalis sp.n. — Clavus proximally without lightish spot; dorsal pubescence dense, silvery; (Mongo- lia): er. kaszabi (Hoberlandt, 1971) 11. Exocorium with narrow ochreous costal margin along entire length (fig. 101, n); (S. Kazakhstan ee kerzhneri sp.n. — Corium with varying light-dark pattern 12 12. Longest semi-erect dark setae of corium longer than diameter of tibia 3 ......... 13 — Longest semi-erect dark setae of corium subequal to or shorter than diameter of tib- TARSIA LOL, LIME AMR Oe 18 13. Pronotum without erect setae extending be- yond lateral margin. Exocorium with only adpressed short, lightish setae, and a sub- marginal longitudinal strip devoid of any setosity; base of exocorium mostly wider than base of endocorium, part of lateral R. H. CoBBEN: Eurasian Saldidae 257 Fig. 20. Teloleuca kuznezovi. a, parameres, specimens from different localities in E. Russia and Japan (second picture from left); b, apex of paramere; c, male grasping plate; d, median endosomal sclerite; left, plane view; right, left lateral view; e, base of penisfilum; f, parandria; g, variation of dark-light pigment pattern of fore wing. 258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 outline of forewing straight (fig. 12c). White-black pattern of corium contrasting (fig. 13h); clavus sometimes with subbasal spot . variabilis connectens (Horváth, 1888) The explanate base of the exocorium of M. variabi- lis is not always clear, and sometimes also occurs in other species, e.g. M. jakovleffi and M. nivalis; these two species, however, lack the hairless exocorial sub- marginal strip. It is still uncertain whether connectens must be considered a subspecies of variabilis or a proper species. I have seen material from Morocco, Yugoslavia, Albania, Bulgaria, Rumania and Greece, corresponding with connectens. True variabilis (endo- corium without erect setae) was seen from Sweden, Belgium, France, Spain, Germany, Poland, Austria, Switzerland, Italy, Hungary, Greece, Iran, Iraq and USSR (Georgia, Armenia). This picture would favour the idea of two separate species, with sympatry in the Balkan countries. However, material seen by me from Turkey and Palestine is more or less intermediate as regards the setosity of the endocorium. — Pronotum with erect setae distinctly ex- tending beyond lateral margin; entire cori- um with erect setae in addition to adpressed Dubescence NPA a 14 14. Erect setae of second tibia longer than the spines; erect setae along entire length of third tibia longer than the spines (see also note under M. scotica, couplet 16 of this key); (Japan, Shikoku) .. shikokuana sp.n. — Setae of second tibia not longer than the Spinesgre Peary ende: 15 15. Erect setae on external proximal part of third tibia longer than the spines ....... 16 — Erect setae on external proximal part of third tibia subequal to or shorter than the SPINEA AA e en ee Ne the 17 16. Lightish pattern of wing predominantly re- stricted to exocorium (fig. 14b); large sub- apical spot on exocorium also persistent in dark specimens; lightish subapical spot of clavus only rarely present. Dorsal pilosity dense; length of erect setae about two times the diameter of the hind tibia; silvery de- cumbent setae dense, semilong, somewhat shagey (STE) MEENDEN e. te fee CERIANO madonica (Seidenstücker, 1961) — Lightish spots distributed on exo- and en- docorium; subapical spot on clavus only absent in dark variants (fig. 14a). Dorsal erect and adpressed pilosity less dense; erect setae subequal to or slightly longer than diameter of hind tibia; (Europe, Euro- pean part of the USSR; earlier records from Japan refer to S. nobilis) scotica (Curtis, 1835) I have seen one male from Turkey (Caycuma, 31.v.1980, leg. Hava) and two females from Caucasus (Krasnaya, no date, leg. Zhelokhovtsev; Azan, 2600 m, above Rhododendron zone, 1.vii.1974, leg. Behäc), with longer pilosity on the legs, nearly as in M. shiko- kuana (couplet 14). In the latter species the dark parts of the wings are ashy-semipruinose black with the distal part of endocorium and medial half of clavus deep satin black. The dark wing pigment of M. scotica is unicolorous black. Additional material is needed for a taxonomic interpretation of this type of geographic variation of M. scotica. 17. Clavus proximally always with small yel- lowish spot near edge bordering the corium and sometimes with an elongate spot in the edge bordering the scutellum (14f). Light- dark pattern of corium contrasting; lightish spots yellowish; spot on mid part of exoco- rium divided longitudinally (fig. 14f). Dark pigment of margin of exocorium and apical area of endocorium intense black, contrast- ing with otherwise ashy black colour; (Ja- DA ier 6 Saree a AR ES miyamotot sp.n. — Proximal part of clavus entirely black. Con- trast between light and dark pattern of cori- um not sharply defined. Light spots greyish white; spot on mid part of exocorium usually large and undivided (fig. 14c). Dark pigmentation uniform ashy-black; (Uzbe- kistan)) eee tadzhika (Kiritshenko, 1912) 18. First acetabula entirely or partly lightish 19 ——| Firstiacetabulalblacky) >. er Veen 21 19. Wings dull, with only adpressed short sil- very setae. Middle trichobothrial-like ce- phalic setae arising nearly always from pale spot. First coxae for major part pale. Wing pattern variable; preponderance of white markings on endocorium; in darker forms markings disappear first on exocorium; (Nlaska) ar... monae (Drake, 1952) — Wings weakly shining; semilong erect setae are present in addition to the short decum- bent coldempubescencel WA 20 20. First acetabula entirely pale. Wing pattern asınetie. 141 (alkwazal: stan) seen VE. Hoa oblonga acetabularis subsp.n. — First acetabula dark with pale apical mar- gin. Corium for major part testaceous (fig. 14k); membrane hyaline. Clavus sometimes with subbasal pale stripe. Legs predomi- nantly yellowish; (Kazakhstan) ........... Lorne ee koktshetavica sp.n. 21. Second antennal segment with only very short pubescence. Proximal part of exocori- um wider than base of endocorium (fig. 12b); part of external outline of left and R. H. COBBEN: Eurasian Saldidae Fig. 21. Salda species scanning micrographs of mid part of corium to show differences in densities of setae (60X). a, S. morio, Japan; b, S. muelleri; c, S. littoralis; d, S. kiritsjenkot. 260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 right wing parallel-sided (not clearly pre- sent in some specimens). Wing dull-black with usually two larger well-demarcated spots on exocorium and some smaller ones on endocorium (fig. 131); stronger tendency to complete melanism than in the subsp. connectens, particularly in specimens from the Caucasus; membrane usually unicolo- rous smoky black. See remarks at couplet 13, sub variabilis connectens........... va- riabilis variabilis (Herrich-Schaeffer, 1835) — Second antennal segment with very short pubescence and with some erect, somewhat longer setae over entire length of segment. Proximal part of exocorium about as wide as base of endocorium or wider; outline of wing margin regular convex ........... 22 22. Erect setae on corium of about same length as the width of third tibia. Colouration of short pubescence of wing mostly golden. Marginal setae along anterior edge of pro- notum shorter than diameter of one ocellus. Wing design usually as in fig. 71, drawings 2 and 3; lightish pattern testaceous, not very contrasting; base of clavus rarely with pale spot along claval suture; membrane smoky with dark patches in between the vernsa(fdiransbatl a me em FAI Be oblonga oblonga (Stal, 1858) — Pubescence of wing clearly shorter than diameter of third tibia. Marginal setae along anterior edge of pronotum longer than or subequal to diameter of one ocellus, rarely shorten cai jakovleffi (Reuter, 1891), mongolica (Kiritshenko, 1912) and nivalis (Lindberg, 1935) (see following discussion). At the end of this key three species remain which caused some nomenclatorial problems: M. jakovleffi, M. mongolica and M. nivalis. | give the following explanation and hope that my proposal on the nomenclature will prove to be correct. M. jakovleffi (Reuter, 1891). Reuter de- scribed this species from Turkestan, Dschiptik, D. Fedtschenko and, judging from his notation “long 2 6—7'2 mill.”, he apparently had the disposal of at least two females. The diagnosis refers to a species without erect setae, with shining head and thorax, and with a number of white spots on the forewing. Reuter pointed out that the species differs from the hirsute M. scoti- ca, but that the differences with M. variabilis are difficult to describe. So far I could not trace the type material of M. jakovleffi, but received one male and one female from the Reuter coll., labelled Turkestan Dschilarik (about 540 km NE of the type locality), leg. J. Sahlberg. The distribution of the lightish wing spots (fig. 14d, second picture from left) comforms more or less with the description of the type material. Since head and pronotum are rather shiny, it seems logical to attribute the male and female in ques- tion provisionally to M. jakovleffi. I propose to designate the male as the neotype in case the original specimens will not emerge!). The two specimens from Dschilarik possess a character not mentioned by Reuter but of possible impor- tance for identification. The laterofrontal edges of the pronotum are beset with a fringe of densely packed whitish setae (fig. 135); the length of these setae is variable in longer series (see below). M. mongolica (Kiritshenko, 1912). I studied one specimen marked with the type indication ‘ of the Leningrad Museum. Since it bears exactly the same locality data as given in the original de- scription (Mongolia: decliv. septentr. Altai Mongolici: ripae lacus Kobdo Inferioris (P. K. Kozlov. viii.1899) I take it for granted that it is the specimen on which Kiritshenko based his description. The body dimensions (5.5. mm— 2.5 mm) also fit the description, but instead of a male, as indicated, it appears to be a female. I provisionally suggested that this single type specimen might be conspecific with M. jakov- leffı as conceived above. The anteromarginal fringe of setae on the pronotum is not well-developed, but as men- tioned before, this character also greatly varies in other material from Mongolia. Additional material from Mongolia seen by me originates from the following locaties: SE Arakhangaj, Sharagoldzi (Shargoldzuty-gol) river, 20—40 km NNE Bajan Khongor, 1 2, 25.v11.1926, leg. Kiritshenko; Ongiin (Ongin)-Gol river, upper part, 50 km NNW Arvaj-Kheer, 5 d 4 9, 14.vii.1926, leg. Kiritshenko (together with M. oblonga; Bajanchonger aimak, Changaj Mts So- mon Zap, river, Zaesol 2100, m 22 Ser 18.v1.1966, expedition Kaszab (Hoberlandt, 1971b, referred to the males (loc. no. 79) as M. oblonga and to the female (log. no. 708) as M. !) Dr Kerzhner informed me very recently that Dr A. V. Sviridov let him know, in a letter of 11.viii.1985, that the type of M. jakovleffi is in the Zoological Museum of the Moscow University; its length is 6.2 mm, the sex is not indicated, and it is labelled: “Dzhiptyk” (in Cyrillic characters), and “Acanthia jakowleffi Reut”. R. H. COBBEN: Eurasian Saldidae 261 mongolica. This material is plotted on map 2, provisionally as M. mongolica, although I ini- tially was inclined to consider them conspecific with M. jakovleffi. After correspondence with Dr Lindskog I now adopt a more conservative attitude. Recent expeditions made in Mongolia and China probably contain large samples of the jakovleffi complex. Scrutinizing this material might solve whether some minor differences re- vealed between true M. jakovleffi and the sparse material of M. mongolica at hand are constant enough to uphold M. mongolica as a valid taxon (species or subspecies). M. nivalis (Lindberg, 1935) (comb. nov.; de- scribed as Acanthia nivalis). The type locality of jakovleffi this species is Kashmir, Tehrong valley near Sia- chen glacier, 4125 m, 20—26.vi.1929, leg. J. A. Sillem, Netherlands Karakorum expedition. The type material located in the Zoological Museum Amsterdam consists of one female and four lar- vae. The second adult mentioned in the original publication could not be traced. Although the single female at hand is in a teneral condition, some characters match the series I could study from mountainous areas north of Kashmir, which differs from M. jakovleffi. Such charac- ters are: dull dorsal cuticle, more dense silvery adpressed setae, and rather porrect anteclypeus (fig. 14g). Despite of the incomplete informa- tion on the type material of M. nivalis I am nivalis size and shape length male length female membrane era colour head and pronotum adpressed setae of corium eunomy of wing longitudinal spot in basal half of endocorium head in front of eyes parandria paramere generally larger, male less slender, female more slender 5.0—6.0 mm (mean 5.4) (n = 13), width 2.0—2.4 mm (mean 2.2); ratio /w 2.0 wu 6.0—7.0 mm (mean 6.4) (n = 11), width 2.6—2.9 mm (mean 2.8); ratio /w 2.3 fully developed black rather shiny usually greyish or golden, not striking fig. 14d undivided in pale specimens, often parallel with adjacent spot of exocorium less porrect (fig. 14h, left) widely incised (fig. 12}) with broad processus sensualis and regular external outline (fig. 12h) 1) Dr Kerzhner wrote me that the variability of wing design is more diversified than presented in fig. 14e. generally smaller, male more slender, female less slender 4.5—5.0 mm (mean 4.7) (n = 13), width 1.9—2.7 mm (mean 2.1); ratio /w 2.24 5.1—6.0 mm (mean 5.5) (n = 13), width 2.3—2.8 mm (mean 2.6); ratio l/w 2.1 er slightly reduced ashy black rather dull silvery and dense (fig. 15a) fig. 14e!) subdivided; marginal spot halfway exocorium (see arrow) mostly present more porrect (fig. 14g, left) due to bulging anteclypeus and more transverse swelling above anteclypeus (fig. 14g, right) incission narrow (fig. 12k) with pointed processus sensualis and somewhat undulated external outline (fig. 12g) 262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 more inclined to treat the material listed below as M. nivalis, rather than to describe it as a new species. Before having studied the type of M. ni- valıs I named this species in 1960, in manu- script, as Saldula horvathi, and I misused this name as nom. nud. in one of the chapters of my book (Cobben 1968: 20) dealing with the eggs of Heteroptera. Making allowance for some mentioned no- menclatorial uncertainties, the differences be- tween M. jakovleffi and M. nivalis can be tabu- lated as above. The material verified by me, mostly present in the Leningrad Coll. and the Coll. Yu. Popov, Moscow, is plotted on map 2; some additional symbols have been added on map 2 in accord- ance with recent information supplied by Dr Kerzhner. M. jakovleffi: Kirghizia, Tien Shan, Kirghizskij Ridge (= Aleksandrovskij Ridge), gorge Kenkol, 4 d 1 2, 17.v11.1930, leg. V. Bianchi; idem, Kirghizskij ridge, Frunzenskaja province (= Semirechenskaja province), 3800 m, 1 ®, 15.vu.1910, leg. Kiritshenko; idem, Terskej Alatau Ridge (southern side of eastern part of Ridge), Kujlu river (100 km SE Issyk-Kul lake), 1 4, 24.v1.1902, leg. Saphozhnikov, idem, Terskej Alatau Ridge, Karagatuz river, 1 d, 26.vi.1902, leg. Sapozh- nikov; idem, Terskej Alatau, Karasaj river (the Upper Naryn river, 80 km S of Issyk-Kul lake), Pokrovskie syrts (plateau), 1 d 2 2, 29.vii.1965, leg. R. Zlotin; idem, Terskej Alatau Ridge, Karasaj river, 1 6, 14.v11.1953, leg. D. Panfilov; idem, Fergansky Ridge (Nw part), Kizylunkur river (the Upper Karaunkur river), 20 km N of Arslanbob, 1 6, 8.vili.1958, leg. Yu. Popov. Syrt Arabel, 1 d 1 ©, no further data, type locality of var. moerens Horv., 1904; Djergalan- Tjuk, leg. Almasy, no further data. Kazakhstan, Tien Shan, Zailysky Alatau Ridge, Bolshaja Almaatinka river (15—20 km S Alma-Ata), 3 9, 25—28.vii.1928, leg. Shnitnikov; idem, Saur Ridge, Karasu river (the Upper Kenderlyk river; 50 km SE Zaisan lake, 9 & 6 Q, 18.vi.1965, leg. Yu. Popov, 3 d 3 9, 20.vi.1965, leg. I. Sukatscheva. Usbekistan, Torino near Tash- kent, 1 2, 13.vii.1924, leg. Martynova; idem, Tshim- gan Mt, Tian-Shan, 2000 m, 5 d, 9.vi.1982, leg. P. Stys. leg. Almäsy. Tadzhikistan, Pamiro-Alaj Alajska- ja valley (eastern part), near Irkeshtam, 1 9, 20.v1.1960, leg. I. Lopatin; idem, Kzyl-Rabat, 1 2, 11.vu.1960, leg. Lopatin; idem, Murgab, 1 4, 24.v1.1937, leg. Luppova & Redikortsev.; Altai, Tsha- gan-Uzun—Taldur, 6.viii.1912, leg. Sushkin; Shavoz, river Shakhdara, 2 d, 10.vi.1956, leg. Zhelokhovtsev. M. nivalis: Type-locality: Kashmir, Tehrong Valley. Kirghizia, Kirghizsky ridge (= Aleksandrovskij Ridge), Fruzenskaja province (= Semirechenskaja province), 3800 m, 1 2, 15.vii.1910, leg. Kiritshenko; idem, Tien Shan, Chatkalskij (Tschatkalskij) Ridge (NE part), Sary-Chilek Lake, 2500 m, 1 d, 13.vin.1957, leg. Yu. Popov; idem, N. Park, Ala-Archa nr. Frunze, 2200 — 2800 m, 14 d 13 @, 22.vi.1982, leg. P. Stys; idem, 2100 m, 1 d 1 2, 6—9.vii.1976, leg. J. Kral; Fergana mer., Alai, Artschi-Bashi, large series, 12.vi.1908, leg. Kiritshenko; Syrt Arabel, leg. Almasy, 2 9, no further data. Kazakhstan, Zailijskij Alatau Ridge, gorge Go- relnyi (15—20 km S Alma-Ata), 2000 m, 1 9, 24.vili.1958, leg. D. Panfilov. It appears from the distribution pattern now known (map 2) that M. nivalis and M. jakovleffi are exclusively mountainous species (registered altitudes from -2000—3800 m with ranges coin- ciding in the Kirghizia area (37—45° latitude). Some locality data and collecting dates are even the same for both species. Dr Kerzhner in- formed me that both were collected near Alma- | Ata in the same valley, but at different sites. The poorly explored mountains at lower latitudes disclosed so far only one locality of M. nivalis (35°) and two females from 27.5° latitude (Bu- than, 20 km S of Thimphu, 2300 m, 18.v.1972, Bhutan expedition Nat. Hist. Mus. Basel) which belong to the jakovleffi-mongolica complex. The range of M. nivalis apparently does not ex- tend to the Mongolian mountain chains, where- as the range of M. jakovleffi seems to continue into W. Mongolia, either with conspecific popu- lations or with a very close relative (M. mongo- lica, see above). COMMENTS ON THE ZOOGEOGRAPHY OF MACROSALDULA, SALDA'), AND TELOLEUCA The ancestors of these three genera apparent- ly originated somewhere in the Palaearctic Re- gion. Only two species of Macrosaldula occur in the Nearctic Region (Alaska), viz. M. rivula- ria Sahlb. and M. monae Dr. The latter species is only known from Alaska, whereas M. rivula- ria is one of the very few Macrosaldula species of which the range extends into the arctic zone (maps 1, 2). The characters of both species, with only one exception, conform with those of Pal- aearctic Macrosaldula. Salda littoralis is the only Holarctic species of the genus. With respect to one phylogenetically most important male geni- tal character which is shared by all Old World species, it is more plesiomorphic than the re- maining seven exclusively New World species. ') Salda henschii Reut. and S. sahlbergi Reut. are left out of account, since Dr P. Lindskog is revising this couple of species. R. H. COBBEN: Eurasian Saldidae 263 Of the genus Teloleuca, the sister group of Sal- da, two of the four Eurasian species also live in the northern part of the New World. This three-fold picture speaks well for a common mi- gration route from the Old World across the Bering Strait after separation of the northern continents. Combining the Eurasian distribution patterns (maps 1—4) the broad band area from Central Europe across Siberia, which is practically with- out any records of the saldids discussed, be- comes apparent. This absence of records cer- tainly coincides with paucity of sampling. The distribution maps of Gerridae (Kanyukova, 1982) reveal a rather similar pattern, although this group of waterstriders seems to have been explored better than shorebugs in the European part of the USSR. It may, however, be expected that even after more extensive exploration of the West Siberian lowlands, the general distribution patterns of the species treated in the present pa- per will not change markedly. The distribution patterns of maps 1—4 make: at first sight clear that species of the three gen- era considered here are predominantly moun- tainous (note the absence of data from the Ural) and that the largest number of species is found in Central Asia, E. Siberia + Mongolia and the Far East (8(3), 11(3) and 9(2) species, respecti- vely; numbers between brackets refer to en- demic species). From Japan seven species are presently known of which two Macrosaldula species seem to be restricted to this archipelago (an additional species from Japan will be de- scribed by Dr J. Polhemus (in litt)). The Euro- pean part of the USSR harbours six to seven species, of which only M. scotica (Curt.) and M. variabilis (H.—S.) are typical European el- ements. Few species are confined to the medi- terranean region: M. heijningeni Cobben and M. madonica Seid. It is too early to discuss the ecological diver- sifications of the various eastern species, be- cause locality data are not detailed enough as re- gards habitat characteristics and altitudinal stra- tifications. Some species are labelled as occurring in the same locality, such as M. inor- nata and M. variabilis, M. variabilis and M. sco- tica, S. muelleri and S. kiritshenkoi, and S. mi- cans, S. splendens and S. morio. Smali-scale eco- logical displacements are predictable in such cases. Of great interest is to know more of the habitat requirements of the atypical Macrosal- dula roborowskü which is only known from two localities (map 1) in W China. Its Chartos- cirta-like facies suggests a habitat which might be quite different from those of its congeners which generally inhabit stony banks along streams. Saldidae, constituting a uniform group of wet-soil-dwelling, carnivorous Heteroptera ex- hibiting a large amount of wing-polymorphism, are as ideal for eco-geographical studies as Car- abidae are among Coleoptea. It is hoped that the present taxonomic contribution will foster large scale collections and detailed field obser- vations of shore bugs in the vast Asian conti- nent. ACKNOWLEDGEMENTS The preparation of this paper would not have been possible without the help of Dr I. Kerzhn- er (Zoological Institute, Leningrad), who spent much time in translating and mapping the Rus- sian locality data. I thank him very much for his continuous interest in this project and for his generosity in sending material all over again. I thank Dr Lindskog and Dr Stys for their com- ments on the manuscript, and Dr Polhemus for permission to list some records of species col- lected by him in Japan in 1980. The assistance of the following colleagues in supplying material is greatly acknowledged: M. Brancucci (Basel), W. R. Dolling (London), S. Drosopoulos (Athens), J. Duffels (Amsterdam), K. K. Gunther (DDR, Berlin), H. Hasagawa (Ibara- ki), E. Heiss (Innsbruck), L. Hoberlandt (Pra- gue), J. Lattin (Corvallis), P. Lindskog (Stock- holm), J. Martens (Mainz), S. Miyamoto (Fuku- oka), R. Poggi (Genoa), J. Polhemus (Englewood), Yu. Popov (Moscow), R. Remane (Marburg), M. Satö (Tokyo), A. Sods (Buda- pest), P. Stys (Prague), M. Tömökuni (Tokyo), T. Vasarhelyi (Budapest). REFERENCES Asahina, S., & H. Hasegawa, 1951. Insects of the Oze District, I. — Oyo-Dobutsu 16: 184—189. Benedek, P., 1970. The semiaquatic Heteroptera in the Carpathian Basin with notes on the distribu- tion and the phenology of the species. — Faun. Abh. 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Die Heteropteren-Ausbeute der Mongolisch-Deutschen Biologischen Expeditio- nen 1962 und 1964. Ergebnisse der Mongolisch- Deutschen Biologischen Expeditionen seit 1962. Nr. 22. — Mitt. Zool. Mus. Berlin 43 (1): 53—76. Wroblewski, A., 1966. Shorebugs (Heteroptera, Saldi- dae) of Poland. — Polskie Pismo Ent. 36 (12): 219—302. Wroblewski, A., 1968. Klucze do oznaczania owadów Polski. Czesé XVIII. Pluskwiaki róz- noskrzydle-Heteroptera. Zeszyt 3 Leptopodidae, Nabrzezkowate-Saldidae. — Polskie Towarzyst- wo Entomologiczne 58: 3—35. Wu, C. F., 1935. Catalogus insectorum sinensium. Vol. II. Hemiptera, etc., 634 pp. — Peiping. Zetterstedt, J. W., 1840. Insecta Lapponica, 1140 cols. — Lipsiae. 266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 AU | Il | i N "li! INNI PAL I | I h h I | NH HR I! | rv m À € | > HU iG MAS ars & HUN HN DI rately | | ul li i HI . ; | | | : i KL) (| i | : | | \ | A HN I; HI il | Ù Hi I | N | a | Ti ut i 7 à | la er K Au > | ms | f ( L T { ‘ a x \ (e ER 267 ian Saldidae AT i Ky Uy ET n I | Hy \ | NT R. H. COBBEN: Euras sy ‘su *ds euenyoyfys “KH =® ‘“qTUBS BFAPTNATI ‘NH = “u "N =)“ PUTTI STFTeapu ‘N =O °;-apy eoFTOZu0m *N =@ ‘vu ‘ds Fozoweiru ‘y =© ezsey *N = ‘‘anoy F3FJOTAONEE N =© ‘cu ‘ds sffeaefo einpfesorsey = À ci ec Mima x | 7 == ri TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 268 Ss Ed 269 tan Saldidae R. H. COBBEN: Euras ‘paddew u90q Jou SAEY puEjo pue ETAEUTPUE “TIea Sueonçred *I =() ‘‘qpurt + 95 UI SAINI[EIOT] ‘sarmunoo sey Ivy pue ISS 241 ur soloads yonajojay zo uonngınsıp UMOUY "+ dew Aozauzny ©] -& ‘-qnoy TXNIZOUCIOQ “I = ‘*sdwoyy eIeTosejtgq eonororor = @ 270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, AFL. 4, 1985 Table 1. Some measurements of Halosalda, Saldula, Macrosaldula, Calacanthia and Salda species (in mm). Data are presented for only four specimens at the most; when longer series are available more data can be found in the descriptions. The length of antennal segments 3 and 4 is inclusive the proximal intersegmental ring, which creates some variation in the measurements since the internodes are not always fully visible. pronotum antenna leg 3 ci > 5 3 = SUS a os È - $ 8 e So = re = D © B og 3 2 | = ge 20 2 33 DE DI 5 CONSE so BElm Se SS So | species DINE 9 Sen 8 8 Sa B B = Halosalda d 34 1.6 0.96 0.32 0.28 0.04 0.07 0.52 0.56 1.16 0.26 0.68 0.44 0.42 1.60 0.64 | coracina sp.n. d 3.6 1.6 0.94 0.36 0.28 0.05 0.05 0.58 0.56 1.20 0.26 0.68 0.44 0.46 1.52 0.68 | 9 3.8 1.9 1.00 0.38 0.32 0.03 0.03 0.54 0.58 1.28 0.28 0.78 0.48 0.44 1.78 0.68 OZ fB 1.00 0.36 0.32 0.03 0.04 0.52 0.60 1.30 0.28 0.76 — — 1.72 0.66 Saldula d 3.05 1.59 0.89 0.34 0.24 0.05 0.05 0.46 0.54 115 0.29 055 0.41 0.41 1.48 0.51 hasegawai S 3.16 1.70 0.95 0.31 0.24 0.05 0.05 0.49 0.55 1.20 0.29 0.57 046 0.45 1.51 0.52 sp.n. 9 355 1.90 0.94 0.35 0.25 0.07 0.04 0.50 0.58 1.30 0.30 0.62 0.45 0.45 1.59 0.56 Saldula d 35 1.75 1.04 0.40 0.26 0.06 0.04 0.48 0.56 1.32 0.36 0.70 — — 1.68 0.64 taiwanensis d 36 1.70 1.04 0.38 0.24 0.06 0.04 0.48 0.56 1.32 0.36 0.68 0.46 0.46 1.64 0.63 | sp.n. 2 4.0 1.90 1.08 0.40 0.32 0.07 0.04 0.52 0.62 1.44 0.38 0.72 — — 1.76 068 - ® 3.75 1.90 1.04 0.40 0.28 0.07 0.04 0.56 0.60 1.40 0.36 0.70 0.45 0.44 1.76 0.63 Saldula d 3.60 1.78 1.08 0.41 0.28 0.07 0.04 0.50 0.65 1.36 0.28 0.70 0.48 0.54 1.75 0.63 burmanica d 4.22 1.99 1.16 0.45 0.32 0.05 0.05 0.60 0.72 1.60 0.29 0.75 0.53 0.58 1.95 0.73 | subsp.n.? ® 414 2.28 1.18 0.48 0.35 0.07 0.05 0.56 0.70 1.63 0.28 0.73 0.50 0.51 1.90 0.75 9% 3.39 210 1.18 0.45 0.34 0.06 0.05 0.56 0.73 1.48 0.28 0.73 0.53 0.57 1.88 0.71 Saldula 6 43. 2:71 1.16 0.35 0.28 0.06 0.03 0.66 0.66 1.35 0.38 0.90 0.57 0.60 2.10 0.85 sibiricola sp.n. SEES 1.20 0.40 0.30 0.06 0.03 0.70 0.70 1.42 0.33 0.90 0.54 0.59 2.15 0.87 9 49 24 1.25 0.42 0.31 0.07 0.03 0.70 0.71 1.55 0.35 1.00 0.62 0.65 2.40 0.96 Macrosaldula d 4.0 1.7 0.96 0.40 0.32 0.08 0.02 0.46 0.56 1.40 0.28 0.76 0.45 0.45 2.20 0.68 clavalis sp.n. ONCE) 7220 1.04 0.44 0.38 0.08 0.03 0.52 0.72 1.64 0.36 0.88 0.60 0.54 2.44 0.78 | Macrosaldula d 43 182 1.0 0.44 0.36 0.08 0.04 0.54 0.64 1.44 0.24 0.80 0.54 0.52 2.10 0.68 | inornata sp.n. d 45 1.8 1.0 0.44 0.36 0.08 0.03 0.54 0.62 1.48 0.24 0.80 0.56 0.54 2.10 0.64 dr 27 «20 1.06 0.48 0.44 0.09 0.04 0.60 0.72 1.72 0.28 0.90 0.58 0.56 2.52 0.76 29 49 20 1.03 0.52 0.40 0.09 0.04 0.52 0.72 1.64 0.27 0.90 0.58 0.54 2.36 0.69 Macrosaldula 6 bo DL 1.28 0.60 0.36 0.08 0.02 0.70 0.76 1.92 0.52 1.40 0.92 0.84 3.24 1.00 kerzhneri DZ 3.0 1.38 0.60 0.48 0.09 0.02 0.80 0.80 2.16 0.56 1.60 0.96 0.88 3.92 1.18 | sp. n. Q 72 3.0 1.40 0.64 0.46 0.09 0.03 0.88 0.84 2.16 0.53 1.56 0.92 0.84 3.84 — Macrosaldula d 4.7 19 1.16 0.48 0.36 0.08 0.02 0.56 0.68 1.56 0.36 0.92 0.54 0.54 2.36 0.88 koktshetavica d 43 1.9 1.14 0.50 0.34 0.07 0.02 056 0.66 1.48 0.36 0.92 0.56 0.54 2.32 0.89 sp.n. 2 50 22 1.20 0.46 0.40 0.09 0.03 0.60 0.72 1.72 0.38 1.02 0.56 0.56 2.48 0.96 2 50. 21 1.20 0.50 0.40 0.08 0.03 0.60 0.68 1.72 0.32 0.96 0.58 0.54 2.40 0.92 Macrosaldula d 4.8 1.9 1.04 0.48 0.32 0.08 0.03 0.48 0.64 1.56 0.32 0.86 0.58 0.56 2.24 0.72 miyamotoi sp.n. d 44 19 0.98 0.38 0.28 0.07 0.03 0.48 0.60 1.52 0.28 0.84 0.46 0.46 2.20 0.76 O Dr 29 1.08 0.44 0.32 0.09 0.04 0.52 0.68 1.68 0.36 0.96 0.64 0.60 2.72 0.78 Macrosaldula d 48 2.0 1.04 0.42 0.30 0.08 0.03 0.46 0.64 1.56 0.32 0.92 0.60 0.58 2.36 0.78 shikokuana sp.n. ON FAT 24 1.08 0.40 0.32 0.09 0.03 0.56 0.66 1.72 2.64 0.76 Macrosaldula d 6.3 2.9 1.36 0.56 0.44 0.08 0.02 0.80 0.80 1.88 0.52 1.60 0.98 0.80 3.76 1.08 simulans sp.n. 2 67 3. 1.32 0.56 0.42 0.12 0.03 1.12 0.80 2.12 0.52 1.52 0.92 0.84 3.60 1.10 9° 65. 29 1.32 0.52 0.43 0.10 0.04 0.76 0.84 2.12 0.56 1.60 0.93 0.85 4.00 1.20 Macrosaldula © SA Dl 1.28 0.48 0.30 0.08 0.04 0.62 0.68 1.80 0.52 1.36 0.64 — 2.56 0.98 violacea sp.n. 6 BA > Bil 1.24 0.48 0.29 0.08 0.02 0.64 0.68 1.72 0.48 1.36 0.72 0.68 2.88 0.96 Japan OBO 22 1.24 0.46 0.36 0.09 0.03 0.64 0.72 1.84 0.48 1.44 0.76 0.72 2.96 1.10 ® 60 2.35 1.24 0.40 0.46 0.09 0.02 0.64 0.72 1.92 0.48 1.32 0.76 — 2.84 1.01 USSR d 5.85 2.12 1.30 0.48 0.34 0.1 0.03 0.65 0.75 1.75 0.55 1.50 0.80 0.72 2.94 1.00 Calacanthia O 75 3.8 1.64 0.78 0.52 0.1 CLO OLR ioe) AL) 07221727 ilo) Ww LG 1026 grandis sp.n. Salda d 60 2.8 1.48 0.49 0.38 0.09 0.04 0.90 0.82 2.05 0.49 1.23 0.78 0.82 3.10 1.25 kiritshenkoi d 54 2.8 1.46 0.42 0.32 0.09 0.04 0.92 0.80 2.00 0.48 1.20 0.80 0.80 2.96 1.20 sp.n. ® 66 3.4 1.72 0.57 0.40 0.10 0.05 1.0 0.98 2.46 0.51 1.34 0.82 0.81 3.55 1.29 semi brachypt. Qu 62 3.4 1.62 0.54 0.43 0.10 0.06 0.92 0.92 2.36 0.50 1.32 0.81 0.81 3.36 1.28 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING REGISTER VAN DEEL 128 * Een sterretje duidt aan een naam nieuw voor de wetenschap * An asterisk denotes a name new to science DIPTERA abietina, Plemiella 206 acariphaga, Ledomyia 200 acerina, Ledomyia 200 Acodiplosis 206, 210 Ametrodiplosis 205 Aphidoletes 200 Arnoldiola 205 Asphondylia 202, 204, 205, [206, 210 Asteromyia 206 Baldratia 205, 206 betulae, Semudobia 208, 209 betulicola, Plemiella 206 Brachineura 196 brachyntera, Thecodiplosis 196. brassicae, Dasineura 195, 203, [204 brevipalpes, Semudobia 209 Buhromyiella 197 cardui, Ledomyia 200 caricis, Antichiridium 205 Camptodiplosis 197 earophila, Lasioptera 203, 204 cerealis, Hybolasioptera 203 Clinodiplosis 197 Contarinia 202, 204 cornifex, Planetella 208 Cystiphora 206 Dasineura 200, 204, 205 deletrix, Rabdophaga 208 destructor, Mayetiola 195, 196, [203, 206 Dichaetia 197 Dichodiplosis 197 Dryomyia 206 Echinella 197 Endaphis 200 Endopsylla 200 Epimyia 196 Feltiella 200 fischeri, Planetella 208 galii, Trotteria 200 galiorum, Schizomyia 206 Gephyraulus 200 Giardomyia 197 Giraudiella 206 Halodiplosis 205, 206 hartigi, Physemocecis 205 Hartigiola 206 heterobia, Rabdophaga 208 hieracii, Cystiphora 206 Hybolasioptera 206 Jaapiella 200, 204, 205 Janetiella 205 Kaltenbachiola 200 Karshomyia 197 kneuckeri, Planetella 208 Lasioptera 204, 205 Lestodiplosis 200 ligustri, Trotteria 200 Loewiola 206, 210 Marcrolabis 200, 204, 205 marginata, Haplodiplosis 195, [203 Mayetiola 205, 206 Mikomyia 208 Misospatha 206 Monobremia 200 Mycetodiplosis 197 Mycocecis 197 Mycodiplosis 197 nasturtii, Contarinia 203 navasiana, Dictyomyia 206 Neolasioptera 204, 205 Neomycodiplosis 197 nervorum, Rabdophaga 208 niveocincta, Lasioptera 206 Oligotrophus 205 oryzae, Orseolia 196 Ozirhincus 206 Phaenobremia 200 pilosellae, Cystiphora 206 pimpinellae, Kiefferia 203, 204 Planetella 205, 206, 208, 210 Polystepha 202 pulchripes, Contarinia 195 Rabdophaga 205, 208, 210 raphanistri, Gephyraulus 203, [204 271 Rhizomyia 196 Rhopalomyia 196, 200, 204, [205, 206, 210 rosaria, Rabdophaga 208 rosenhaueri, Planetella 208 saliciperda, Rabdophaga 208 salsolae, Dictyomyia 206 sarothamni, Asphondylia 206 Semudobia 200, 208, 209 sisymbrii, Dasineura 203, 204 skuhravae, Semudobia 208 steenisi, Semudobia 208, 209 Stefaniola 196, 205, 206 striatum, Antichiridium 205 subterranea, Planetella 208 tami, Schizomyia 206 tarda, Planetella 208 tarda, Semudobia 208, 209 terminalis, Rabdophaga 208 Therodiplosis 200 triandraperda, Rabdophaga 208 tumorifica, Planetella 208 ulmi, Physemocecis 205 Wachtliella 200, 204, 205 HETEROPTERA adriatica, Salda 253, 254 andrei, Saldula 232 angulosa, Calacanthia 242 azteca, Saldula 232 bouchervillei, Saldula 221, 226 brancziki, Teloleuca 252 buenoi, Salda 246 burmanica, Saldula 223 sqq Chartoscirta 215, 223 cincta, Chartoscirta 223 *clavalis, Macrosaldula 215, 219, [229, 256 concolor, Halosalda 218 sqq *Coracina, Halosalda 215, [217 sqq dilutipennis, Chartoscirta 256 fletcheri, Saldula 223 *grandis, Calacanthia 215 *hasegawai, Saldula 215, 221 272 heijningeni, Macrosaldula 255 illinoiensis, Saldula 226 inoana, Saldula 223 *inornata, Macrosaldula 215, [229, 255, 256, 263 jakovleffi, Macrosaldula 215, [232, 260, 262 kaszabi, Macrosaldula 229, 256 *kerzhneri, Macrosaldula 215, [232, 256 *kiritshenkoi, Salda 215, 244 sqq, : [263 *koktshetavica, Macrosaldula [215, 234 koreana, Saldula 232, 240, 256 lateralis, Halosalda 218 sqq littoralis, Salda 215, 246, 262 ssp. piechockii 215, 252 madonica, Macrosaldula 232, [258 micans, Salda 215, 247, 250, [263 Micracanthia 215 *miyamotoi, Macrosaldula 215, [234, 258 monae, Macrosaldula 238, 258, [262 mongolica, Macrosaldula 215, [232, 260 morio, Salda 246, 247 sqq, 250 muelleri, Salda 247, 249, 263 nevadensis, Salda 215, 252 nivalis, Macrosaldula 215, [260 sqq nobilis, Saldula 228 oblonga, Macrosaldula 215, 232, [260 *ssp. acetabularis 236, 250 orbiculata, Saldula 221 orthochila, Saldula 225 pellucens, Teloleuca 228 reuteri, Saldula 228 rivularia, Macrosaldula 232, [238, 256, 262 roborowskii, Macrosaldula 215, [254, 256, 263 saltatoria, Saldula 226 scotica, Macrosaldula 232, 258, [263 *shikokuana, Macrosaldula 215, [236, 258 *sibiricola, Saldula 215, 226 *simulans, Macrosaldula 215, 238 splendens, Salda 215, 247, 263 tadzhika, Macrosaldula 232, 258 “taiwanensis, Saldula 215, 221 tibetana, Calacanthia 242 uichancoi, Saldula 223 variabilis, Macrosaldula 232, [256, 263 ssp. connectens 258 *violacea, Macrosaldula 215, 238, [256 HOMOPTERA Arfaka 165 *breddini, Prasia 165, 166, 169, [174 culta, Prasia 165, 166, 171 Drepanopsaltria 166, 167 faticina, Prasia 165, 166, 169, [170 foliata, Prasia 165 fulva, Arfaka 166 hariola, Arfaka 165, 166 Iruana 165 Jacatra 165 Lacetas 165 Lembeja 165 *nigropercula, Prasia 165, 166, [169, 180 Prasia 165 sqq princeps, Prasia 165, 166, 169, [182 Sapantanga 165 *sarasinorum, Prasia 165, 166, [169, 176 *senilirata, Prasia 165, 166, 169, [186 tincta, Lembeja 165, 166 *tuberculata, Prasia 165, [166, 169, 178 LEPIDOPTERA Acalyptris 8, 45 admiranda, Ectoedemia 18 aegilopidella, Ectoedemia 12, [14, 17, 38, 42, 43, 63, 88 agrimomella, Nepticula 66 agrimoniae, Ectoedemia 6, [12-14, 17, 27, 64, 66 sqq, 69, [71, 87 agrimoniae, Nepticula 66 agrımoniella, Nepticula 66 albifasciella, Ectoedemia 16, 36, [52 sqq, 55, 56, 58, 59, 91 albifasciella, Nepticula 52 albifasciella complex, [Ectoedemia 11, 14, 34, 37, 47, [51, 52 sqq, 64, 88, 91 *algeriensis, Ectoedemia 1, 11, [16, 43 sqq, 47, 88, 90, 91 cf algeriensis, Ectoedemia 15, [43, 45 alliatae, Nepticula 49 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, 1985 *alnifoliae, Ectoedemia 1, 10, 16, [50, 51 amanı, Ectoedemia 10, 13, 15, [18, 24-26, 86, 87 *andalusiae, Ectoedemia 1, 12, [15, 16, 41, 42, 47, 90 angulifasciella, Ectoedemia 12, [14, 69 sqq, 72-74, 77 angulifasciella, Nepticula 69 angulifasciella complex, [Ectoedemia 17, 63, 64, 67, 69, (75, 91 angulifasciella group, [Ectoedemia 2, 63 sqq, 90, 91 apicella, Nepticula 35, 36, 53 arcuata, Nepticula 73 ‘arcuatella, Ectoedemia 12-14, [69, 71, 73-75 arcuatella, Nepticula 73 arcuosella, Nepticula 73 argentipedella, Ectoedemia 79 argentipedella, Lyonetia 78 argentipedella, Nepticula 80 argyropeza, Ectoedemia 12, 33, [34, 35 sqq, 53, 88 argyropeza, Lyonetia 35 argyropeza, Nepticula 31 argyropeza sensu Stainton, [Nepticula 52 argyropezella Herrich-Schäffer, [Nepticula 31 argyropezella Doubleday, [Nepticula 35 aterrima, Nepticula 71, 72 aterrimoides, Nepticula 71, 72 atricolella, Nepticula 71 atricollis, Ectoedemia 12-14, 69, [71 sqq, 75, 76 atricollis, Nepticula 71 atrifrontella, Ectoedemia 10, 13, [15, 18 sqq, 21, 22, 24, 86 atrifrontella, Trifurcula 18 bistrimaculella, Nepticula 82 Bohemannia 8 brunniella, Nepticula 69, 70 canutus, Ectoedemia 28 caradjai, Ectoedemia 7, 11, 15, [16, 38 sqq, 40, 46, 58 caradjai, Nepticula 38 castaneae, Ectoedemia 18 castaneae group, Ectoedemia 17 catharticella, Stigmella 48 cerris, Ectoedemia 16, 52, 54, 55 cerris, Nepticula 54 chasanella, Ectoedemia 38 *contorta, Ectoedemia 1, 16, 151553, 55, 56.9192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, 1985 cursoriella, Nepticula 57, 58 Dechtiria 2, 27 Ectoedemia 1 sqq, 27 sqq, [87 sqq erythrogenella, Ectoedemia 12, [15, 16, 52, 64, 65, 87, 90-92 erythrogenella, Nepticula 64 Etainia 6, 8, 86 euphorbiella, Nepticula 39 Fomoria 1, 6, 8, 10, 29, 86 gilvella, Nepticula 82 gilvipennella, Ectoedemia 8, 10, [15, 16, 43, 45, 46, 55, 62, 88, 90 gilvipennella, Stigmella 45 groschkei, Nepticula 39 hannoverella, Ectoedemia 11, [14, 17, 30-33 hannoverella, Nepticula 30 haraldi, Ectoedemia 11, 15, 16, [41, 44, 47-49, 52, 91 haraldi, Nepticula 47 heringella, Ectoedemia 6, 10, 15, [16, 29, 42, 44, 45, 47-50, 91, 92 heringella, Nepticula 49 heringi, Ectoedemia 53, 56, 57, [59 sq heringi, Nepticula 59 heringiella, Zimmermannia 18 hexapetalae, Ectoedemia 12, 14, [17, 63, 67-69, 77, 87, 91 hexapetalae, Nepticula 68 *hispanica, Ectoedemia 1, 10, 13, [21-23 houzeaui, Nepticula 35 ilicella, Nepticula 47 ilicis, Ectoedemia 10, 15, 16, 29, [44, 45, 47-50, 91, 92 ilicis, Nepticula 48 intimella, Ectoedemia 5, 10, 14, [16, 28 sq, 43, 48, 88, 90 intimella, Nepticula 28 juncta, Stigmella 64 klimeschi, Ectoedemia 7, 12, 14, [17, 32, 33 sqq, 36, 88 klimeschi, Nepticula 33 ladaniphila, Parafomoria 49 Laqueus 1, 8, 86 *leucothorax, Ectoedemia 1, 11, [15, 16, 38, 43, 46, 47, 90, 91 liebwerdella, Ectoedemia 5, 10, 315171872072 192466 liechtensteini, Ectoedemia [55-57, 59, 61, 62 liechtensteini, Nepticula 61 liguricella, Ectoedemia 10, 13, [15, 25 sqq, 86 lindquisti, Ectoedemia 78, 79 longicaudella, Ectoedemia 10, [13, 15, 18-23, 86, 87 mahalebella, Ectoedemia 12, 15, [17, 63, 67-69, 71, 75, 77, 78, 91 mahalebella, Nepticula 77 malivora, Nepticula 71, 72 marionella, Stigmella 31 mediofasciella, Ectoedemia 80, [81 mediofasciella, Tinea 78 minimella, Acalyptris 8 minimella, Ectoedemia 11, 14, [17, 78-80 sqq minimella, Elachista 80 minorella, Nepticula 69, 70 *monemvasiae, Ectoedemia 1, 7, [10, 13, 15, 21, 23, 24, 87 montissancti, Nepticula 54, 55 morosella, Nepticula 35 mucidella, Tinea 78 Nepticula 2 niculescui, Stigmella 33, 35 Niepeltia 8 nigrociliella, Microsetia 57, 58 nigrosparsella, Ectoedemia 8, [10, 14, 16, 50-52, 90, 91! nigrosparsella, Nepticula 51 *nuristanica, Ectoedemia 1, 13, [15, 25, 87 occultella, Ectoedemia 11, 14, [17, 78 sqq occultella, Phalaena 78 occultella group, Ectoedemia [27, 78 sqq, 91 Parafomoria 8 peiuu, Stigmella 21, 22 Phyllocnystis 88 phyllotomella, Ectoedemia 56, [59, 62 phyllotomella, Stigmella 62 populella, Ectoedemia 27, 28 populella group, Ectoedemia [28 sq, 43, 87, 88 populi-albae, Nepticula 31, 33 preisseckeri, Ectoedemia 11, 14, [16, 37, 38, 47, 52, 87, 88, 90 preisseckeri, Nepticula 37 preisseckeri group, Ectoedemia [37 prinophyllella, Nepticula 48 prinophyllella, Stigmella 47 prunivora, Nepticula 71 pubescivora, Ectoedemia 16, [52-56, 92 pubescivora, Nepticula 55 quercifoliae, Ectoedemia 56, 57 quercifoliae, Nepticula 59, 60 278 quinquella, Ectoedemia 11, 15, [16, 43-45, 90-92 quinquella, Microsetia 43 rubifoliella, Ectoedemia 64 rubivora, Ectoedemia 12, 14, [71-75 rubivora, Nepticula 74 rubivora sensu Walsingham, [Nepticula 64 sativella, Ectoedemia 56, 57, 60 sativella, Nepticula 59 schleichiella, Ectoedemia 69 schleichiella, Nepticula 69, 70 septembrella, Fomoria 48 simplicella, Nepticula 35, 36 sivickisi, Ectoedemia 18 species (specimen 1375), [Ectoedemia 62 species (specimen 1843), [Ectoedemia 39 spinosella, Ectoedemia 12, 13, [15, 17, 63, 71-77, 91 spinosella, Nepticula 75 spiraeae, Ectoedemia 7, 12, 14, [17, 64-71, 87-91 spireae, Stigmella 65, 66 staphyleae, Ectoedemia 69 staphyleae, Nepticula 71, 72 Stigmella 2, 29, 78, 79, 88 strigilella, Tinea 78 subapicella, Nepticula 52, 53 subbimaculella, Ectoedemia 39, [52, 53, 56, 57 sqq, 62 subbimaculella, Tinea 57 subbimaculella complex, [Ectoedemia 11, 14, 16, 56 sqq, [82,91 subbimaculella group, [Ectoedemia 43 sqq, 87, 90, 91 suberis, Ectoedemia 12, 15, 16, [34, 38, 40 sqq, 47, 49 suberis, Nepticula 40 suberis group, Ectoedemia [38 sqq, 88, 91 terebinthivora, Ectoedemia 6, [12, 14, 16, 27, 42, 63, 87, 90 terebinthivora, Trifurcula 63 terebinthivora group, [Ectoedemia 63 Trifurcula 2, 8, 10, 26, 45 turbidella, Ectoedemia 6, 11, 14, [16, 30, 31 sqq, 36, 88 turbidella Zeller, Nepticula 31 turbidella Herrich-Schaffer, [Nepticula 35 turbulentella, Nepticula 35 ulmella, Ectoedemia 88 274 utensis, Nepticula 68-70 viridella, Nepticula 40, 41 viridicola, Nepticula 80 weaveri, Ectoedemia 81 wilkinsoni, Ectoedemia 28 woolhopiella, Ectoedemia 79, woolhopiella, Nepticula 80 [81 zimmermanni, Ectoedemia 56, [57, 60 Brassica campestris 195 Brassica juncea 195 Fagus 18, 208 Fagus sylvatica 20 Filipendula 69 Filipendula hexapetala 68 Filipendula vulgaris 68, 70 Fragaria moschata 73 Fragaria vesca 73 Galium 206 Haloxylon 196, 203, 206 Holcus 206 Populus x canadensis 31 Potentilla erecta 73 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 128, 1985 Quercus frainetto 38 Quercus ilex 18, 25, 40, 47, 49, [50 Quercus infectoria 38, 39 Quercus macrolepis 42 Quercus petraea 38, 44, (51-53, 56-58, 60, 61 Quercus pubescens 19, 38, 39, [51-53, 55-58, 60, 61 Quercus pyrenaica 58 zimmermanni, Nepticula 59 Humulus 205 Quercus robur 19, 22, 44, 52, Zimmermannia 1 sqq, 17 sqq, Hypericum 204 [53, 56-58, 60 [86,87 Indocarex 208 Quercus rotundifolia 40, 44, 47, Inula 206 [49 PLANTAE Lathyrus 203 Quercus rubra 58 Abies 206 Lens 204 Quercus suber 23, 40, 46, 47, 49 Aegilops 206 Malus sylvestris 72 "Rosa sempervirens 70 Agrimonia eupatoria 67 Medicago 204 Rosa 70 Agropyron 206 Melilotus 204 Rubus arcticus 74 Alnus viridis 81 Ostrya 88 Rubus caesius 74 Archieracium 206 Phalaris 206 Rubus chamaemorus 74 Aremonia agrimonoides 67 Phleum 206 Rubus fruticosus 64, 74 Artemisia 210 Phragmites 206 Rubus idaeus 74 Artemisia tridentata 196 Pilosella 206 Rubus saxatilis 74 Avena 206 Pistacia terebinthus 63 Rubus ulmifolius 64 Avena sativa 195 Pisum 204 Salix 88, 208 Benzoin 205 Poa 206 Salix caprea 29 Betula 79, 81, 82, 206, 208 Populus 28, 88, 208 Salix cinerea 29 Betula nana 81 Populus alba 32, 34 Salix fragilis 29 Betula pendula 81 Populus canescens 32 Salix pentandra 29, 79 Betula pubescens 81 Populus nigra 31 Salix phylicifolia 29 Brachypodium 206 Populus tremula 36 Salsola 206 Sanguisorba minor 70 Sanguisorba officinalis 70 Brassica napus 195 Potentilla sterilis 73 Santolina 206 Brassica nigra 195 Primocarex 208 Sarothamnus 19, 206 Calamagrostis 206 Prunus avium 72, 77 Secale 206 Capparis 208 Prunus cerasifera 72, 76 Senecio 203 Capsicum 204 Prunus cerasus 77 Serratula 206 Carduus 208 Prunus cocomilia 77 Sesamum 204 Carex 205, 206, 208 Prunus domestica 76 Sinapis 204 Carpinus 18 Prunus dulcis 76 Solanum 204 Castanea 18, 22, 23, 53, 56, 57 Prunus fruticosa 76, 77 Sonchus 203 Castanea sativa 56, 60 Prunus mahaleb 72, 77 Spiraea japonica 66 Centaurea 206 Prunus spinosa 72, 76 Spiraea media 66 Ceratonia siliqua 204 Prunus tenella 77 Spiraea salicifolia 66 Chaematia 208 Pyrus communis 72 Staphylea pinnata 72 Cirsium 203 Quercus 7, 18, 26, 38, 43, 46, Tilia 205 7 Clematis 205 [88, 90, 91, 205, 206 Ulmus 24, 37, 205 Corylus 208 Quercus alnifolia 50, 51 Urginea 204 Corylus avellana 81 Quercus baloot 25 Verbascum 204 Crataegus 72 Quercus cerris 46, 52, 55-58, Vicia 204 Cynodon 206 [61, 62 Dactylus 206 Quercus coccifera 18, 26, 38-40, Ephedra 205 [42, 46, 47 Eruca 204 Quercus faginea 19, 40, 60 y à} # ta ue n DEEL 129 1986 TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING OLOG Tijdschrift voor Entomologie, deel 129, 1986 NEDERLANDSE ENTOMOLOGISCHE VERENIGING BESTUUR (BOARD) Voorzitter (Chairman) 0300000 avvio C. A. W. Jeekel Vice-voorzitter (Vice-President) ................. L. H. M. Blommers Secretanis(Secrctan) PRE OC R. de Jong AGISCE a ae Ryksmuseum van Natuurlyke Historie, Raamsteeg 2, Leiden 2311 PL lesRenningmeestet @ireasuter |) ie sen L. P. S. van der Geest PER SR ee TS al RL AE AUS ga ee Doornenburg 9, Landsmeer 1211 GP 2e Penningmeester (Treasurer Il). .<..:.5..-°..- A. P.J. A. Teunissen CAAT ES GRR E ALE ACEA Strausslaan 6, Vlijmen 5251 HG Bibliothecanisn(sibraniam) incr yore W.N. Ellis VAIN GER ER ne Plantage Middenlaan 64, Amsterdam 1018 DH eid (Member terse haere ern B. van Aartsen TIJDSCHRIFT VOOR ENTOMOLOGIE Redactie (Editorial Board) . ..................... P. J. van Helsdingen, C. van Achterberg, S. A. Ulenberg, J. van Tol, E. van Nieukerken ACCISE RE RO ANTI Ryksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden 2311 PL The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea. Subscription rate: D.Fl. 300,— per year. Issues 1—4 appeared on 10.XI.1986 Issues 5—7 appeared on 15.X11.1986 Issues 8 and 9 appeared on 31.VII.1987 ISSN 0040-7496 INHOUD Bolland, H. R. — Review of the systematics of the family Camerobiidae (Acari, Raphignathoidea). I. The genera Camerobia, Decaphyllobius, Tillansobius, and Tycherobius............ Davis, A. J. — Bibliography of the Ixodiphagini (Hymenoptera, Chalcidoidea, Encyrtidae), parasites Omnicksi (Alcan slxodidae)s withimoteston thei biolooyae lnm |e RO Hensen, R. V. — Revision of the subgenus Prosceliphron Van der Vecht (Hymenoptera, Sphecidae) Jong, M. R. de. — Taxonomy and biogeography of Oriental Prasiini. 2. The foliata group of the genus KombejaDistant 1892 (Homopteran ibienidao ne Lieftinck, M. A. — New and little known Platycnemididae and Coenagrionidae from New Guinea and Tie Sommo Islam (Coma) er ee on Oosterbroek, P., zie Theowald, Br. Pape, T. — A phylogenetic analysis of the Woodlouse-flies (Diptera, Rhinophoridae) ............ Pfau, H. K. — Untersuchungen zur Konstruktion, Funktion und Evolution des Flugapparates der BibéllénitinsecratOdonara) este LEI RR Sota Ae Roskam, J. C. — Biosystematics of insects living in female birch catkins, IV. Egg-larval parasitoids of the genera Platygaster Latreille and Metaclisis Forster (Hymenoptera, Platygastridae) Theowald, Br. & P. Oosterbroek. — Zur Zoogeographie der westpalaearktischen Tipuliden. VII. Die Mipulidentder BalkanhalbinsellDipteralupulidae) nr ae eee: Tol, J. van, zie Lieftinck, M. A. 125 Tr 5 en x uo vind she di N 0 bi oc Le aes ae pe ad te DEEL 129 AFLEVERING 1 1986 JK 6) & | TIJDSCHRIFT VOOR ENTOMOLOGIE f u UITGEGEVEN DOOR f DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING, _ INHOUD Br. THEOWALD und P. OosTERBROEK. — Zur Zoogeographie der Westpalaeark- tischen Tipuliden. VII. Die Tipuliden der Balkanhalbinsel (Diptera, Tipulidae), bid li Tijdschrift voor Entomologie, deel 129, afl. 1 Gepubliceerd 10-XI-1986 Sa ak did vr n° ù ZUR ZOOGEOGRAPHIE DER WESTPALAEARKTISCHEN TIPULIDEN. VII. DIE TIPULIDEN DER BALKANHALBINSEL (DIPTERA, TIPULIDAE) von BR. THEOWALD und P. OOSTERBROEK Zoölogisch Museum (Entomologie), Amsterdam EINLEITUNG Unter Balkanhalbinsel (Balkan) sind in dieser Arbeit nachfolgende Staaten zusammengefaßt: Jugoslawien, Rumänien, Albanien, Bulgarien, Griechenland und die europäische Türkei. Von Teilgebieten der Balkanhalbinsel wurden schon in der zweiten Hälfte des 19. Jahrhunderts Ti- puliden beschrieben und Artenlisten erstellt: Kowarz (1873), Strobl (1897, 1900, 1904), Thal- hammer (1900). Aber erst in der zweiten Hälfte des 20. Jahrhunderts wurde die Tipulidenfauna dieser Halbinsel eingehender studiert. Simova (ab 1959) veröffentlichte über die Tipuliden von Jugoslawien und Erhan (ab 1959) über die von Rumänien. Die Tipulidenfauna von Griechen- land wurde bekannt durch Mannheims (1954) und Theischinger (ab 1977). Über Tipuliden von Albanıen wurde durch Mannheims (1966), über die von Bulgarien durch Szilady (1934) be- richtet. Ziemlich umfangreiches Material von vielen Fundorten auf der Balkanhalbinsel, über das bis heute noch nicht veröffentlicht wurde, findet sich in den zoologischen Museen von Amsterdam, Bonn und London. Aufgrund von Veröffentlichungen und Sammlungen konnten wir eine Liste mit insge- samt 201 Arten zusammenstellen, die mit Si- cherheit von der Balkanhalbinsel nachgewiesen sind. Diese Arten werden in Tabellen erfaßt und analysiert. In diesen Tabellen sind die größeren Balkanstaaten unterteilt (Karte 1). Für Jugosla- wien wurde die Einteilung in Republiken be- nutzt, wie Simova sie in ihren Arbeiten ver- zeichnet. Rumänien wurde aufgeteilt in Südru- mänıen (Walachei, Südkarpaten und Banat- region), in Dobrudscha (mit Donaudelta) und in Nordrumänien (Ostkarpaten und Siebenbür- gen). Griechenland ließ sich am besten einteilen in Südgriechenland (Peloponnes), Mittelgrie- chenland (nach dem Norden bis zum Fluß Pi- nıon), Nordgriechenland (das griechische Ma- zedonien nach dem Osten bis Thessaloniki) und Nordostgriechenland (von Thessaloniki nach dem Osten bis zur europäischen Türkei). Von Nordostgriechenland und von der europäischen Türkei sind insgesamt weniger als zehn Arten bekannt, weshalb diese beiden Gebiete nicht in die Tabellen aufgenommen wurden. Herr G. Theischinger war so freundlich, das Manuskript kritisch zu lesen und sprachlich zu korrigieren. Ihm sei herzlich gedankt. TABELLEN Die 201 Arten der Balkanhalbinsel sind in ae Tabellen erfaßt: . die Arten der europäischen Tiefebenen und die mit ihnen nächstverwandten Arten mit rein balkanischer Verbreitung; 2. die Arten der europäischen Gebirge und die mit ıhnen nächstverwandten Arten mit rein balkanischer Verbreitung; 3. mediterrane Arten. In der letzten Spalte jeder Tabelle sind mit Buchstaben Bone uncer gegeben. Es be- deutet: A. rezent ausgewandert nach Italien, Iberien und/oder Kleinasien; E. endemische Art, aber mit nächstverwandter Schwesterart in Mittel- und/oder Westeu- ropa; rezent eingewandert aus Italien oder Iberien; . rezent eingewandert aus Kleinasien; . ausgewandert nur nach Osteuropa; . rezent eingewandert aus Zentralasien. Tabelle 4 gibt eine detaillierte Zusammenfas- sung der Tabellen 1—3. NOR Tabelle 1: Arten der europäischen Tiefebenen Nach dem Saalien haben sich viele Arten von der Balkanhalbinsel bis in die Laubwalder und Wiesen von Mittel-, West- und Osteuropa aus- gebreitet. Sie wurden im Weichselien haupt- 2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 1, 1986 Tabelle 1. Tipuliden-Arten der europaischen Tiefebenen. (Unter Bemerkungen: E endemische Art, aber mit nachstverwandter Schwesterart in Mittel- und/oder Westeuropa; I rezent eingewandert aus Italien oder Iberien; O ausgewandert nur nach Osteuropa; Z rezent eingewandert aus Zentralasien.) È CMS i a È È 5 Ÿ + Ÿ © (S| so bb ‘E = E è Se È 5 5 5 EE EE Sok SY OS as MIE HD Sa} © = © SEE me) = © a | oO := 3 3 © :3 oO % on Mf) ey re A CZ ra Ctenophora elegans Meigen, 1818 + * PIE fastuosa (Loew, 1871) + + + + Z festiva Meigen, 1804 + + de dp + + flaveolata (Fabricius, 1794) + + + + + + guttata Meigen, 1818 de de 4 + + de 4 ornata Wiedemann, 1818 dp xp Gp PAR Abi aR oF de dp pectinicornis (Linnaeus, 1758) Ar de + Dictenidia bimaculata (Linnaeus, 1758) er = + + + + + + Nephrotoma aculeata (Loew, 1871) + + dp + ab + + analis (Schummel, 1833) + de + + + appendiculata (Pierre, 1919) qe Oar de SR de Se SP Sp de SP dp sp + cornicina (Linnaeus, 1758) ae SP de Se + + de dp croceiventris lindneri (Mannheims, 1951) dd AP an HER ELSE ran crocata (Linnaeus, 1758) de dede + dp ap + + dorsalis (Fabricius, 1781) ae nap sh + + flavescens (Linnaeus, 1758) +.+ + + + + + + I flavipalpis (Meigen, 1830) + + + + | guestfalica (Westhoff, 1880) + HUIT + lunulicornis (Schummel, 1833) + ‘e + oa pratensis (Linnaeus, 1758) dede + + ap + quadrifaria (Meigen, 1804) qe + + + d + + + ie + quadristriata (Schummel, 1833) + + ++ scalaris (Meigen, 1818) + + + + + + + + ++ + + scurra (Meigen, 1818) + = sh + submaculosa Edwards, 1928 + + I Nigrotipula nigra (Linnaeus, 1758) + + + Æ + + + Tanyptera atrata (Linnaeus, 1758) PP + + ap an ae de + nigricornis (Meigen, 1818) > ar Ar Tipula (Acutipula) fulvipennis De Geer, 1776 NEE + + + + + luna Westhoff, 1879 dede HE + À + Se + maxima balcanica Vermoolen, 1983 + + + + + + + + + + + E tenuicornis Schummel, 1833 + + + + + O vittata Meigen, 1804 SE Se ar AP ar SP I (Beringotipula) unca Wiedemann, 1817 + + + + (Dendrotipula) flavolineata Meigen, 1804 + + + + (Lunatipula) fascipennis Meigen, 1818 den dp de op + + dede SE helvola Loew, 1873 ae dp Se dp AP aps dp + + it livida Van der Wulp, 1858 + + + + + + + + + lunata Linnaeus, 1758 HEISSE + + +++ + mellea Schummel, 1833 + + + O THEOWALD & OOSTERBROEK: Westpalaearktischen Tipuliden. VII 3 (Tabelle 1, Fortsetzung) peliostigma Schummel, 1833 selene Meigen, 1830 stubbsi Theischinger, 1979 vernalis Meigen, 1804 (Odonatisca) nodicornis Meigen, 1818 (Platytipula) luteipennis Meigen, 1830 (Pterelachisus) irrorata Macquart, 1826 pabulina Meigen, 1818 pseudovariupennis Czizek, 1912 submarmorata Schummel, 1833 truncorum Meigen, 1830 varipennis Meigen, 1818 (Savtshenkia) alpium Bergroth, 1888 obsoleta Meigen, 1818 rufina Meigen, 1818 (Schummelia) varucornis Schummel, 1833 (Tipula) oleracea Linnaeus, 1758 paludosa Meigen, 1830 subcunctans Alexander, 1920 (Vestiplex) hortorum Linnaeus, 1758 nubeculosa Meigen, 1804 scripta Meigen, 1830 (Yamatotipula) caesia Schummel, 1833 couckei Tonnoir, 1921 decipiens Czizek, 1912 lateralis Meigen, 1818 latemarginata coerulescens Lackschewitz, 1923 marginella Theowald, 1980 montium Egger, 1863 pruinosa Wiedemann, 1817 pierre: Tonnoir, 1921 submontium Theowald & Oosterbroek, 1981 sächlich in das Balkanrefugium, aber auch nach Italien und Iberien zurückgedrängt. Arten, die das Weichselien in mehreren Refugien ver- brachten, haben sich manchmal in heute allopa- trisch vorkommende Schwestertaxa aufgeteilt. Nach dem Weichselien kamen vor allem die Ar- ten des Balkanrefugiums wieder nach dem Nor- den züruck (Theowald & Oosterbroek, 1983). Insgesamt 72 der 201 Arten der Balkanhalb- Slowenien +++ è el N o = Ww ° CS -— oO OMR 1 le Gi 0} fet SE Boe wo) a ee =| OS Fi Boos te GF A a fs 200 ECC EE NE Ene are SR ok SD D © Ss à D © o 30 ESSEC SH SSE SE A) A ey Ser NZ + + + + + + + + + 4+ + + + E + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + - + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + ME EE SM + + + + + + + + + + + + + + + + + - + + + + + + + + + + + + + + + + + + + + - + + + + + + Pate ©) +++ + + + + + + + + + + + + + + + + + + + + + + + + + + insel (36%) gehören zu dieser Gruppe. Die meisten (61) haben sich nach dem Weichselien unverändert bis nach Mittel- und Westeuropa ausgebreitet, drei (in Tabelle 1 mit O gedeutet) nur nach Osteuropa. Insgesamt fünf haben sich nach dem Weichselien von Iberien und/oder Italien bis auf die Balkanhalbinsel ausgebreitet (Tabelle 1 mit I) und eine (Ctenophora fastuosa) breitete sich von Ost- und Zentralasien nach 4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 1, 1986 Tabelle 2. Tipuliden-Arten der europaischen Gebirgen. (Unter Bemerkungen: E endemische Art, aber mit nachstverwandter Schwesterart in Mittel- und/oder Westeuropa.) Slowenien Kroatien Bosn. u. Herz. Serbien Montenegro Mazedonien Albanien Nordgriech. Mittelgriech. Südgriech. Bulgarien Dobrudscha Südrumänien Nordrumänien Bemerkungen Dolichopeza fuscipes Bergroth, 1889 - graeca Mannheims, 1954 + Nephrotoma tenuipes (Riedel, 1910) hate vate ar ARME Tipula (Emodotipula) “saginata” Bergroth, 1891 dr SAP + + =F (Lunatipula) alpina Loew, 1873 al circumdata Siebke, 1863 + + fascingulata Mannheims, 1966 + +++ + + + laetabilis Schummel, 1833 + SF limitata Schummel, 1833 (Mediotipula) sarajevensis Strobl, 1900 + siebkei Zetterstedt, 1852 de Sp + stigmatella Schummel, 1833 de de dE dd dp de dee dk + + (Pterelachisus) austriaca (Pokorny, 1887) + ar crassiventris Riedel, 1864 + + + + + + glacialis (Pokorny, 1887) + +P luridorostris Schummel, 1833 ate eat mayerdueri Egger, 1863 neurotica Mannheims, 1966 + plitviciensis Simova, 1962 pseudoirrorata Goethgebuer, 1921 + pseudopruinosa Strobl, 1895 + (Savtshenkia) aspromontensis Theowald, 1973 benesignata Mannheims, 1954 + + cheethami Edwards, 1924 eleonorae Theischinger, 1978 gimmerthali Lackschewitz, 1925 goriziensis Strobl, 1893 ih + grisescens Zetterstedt, 1838 sp

AE el AS CZ E Be lees e ee 2— 1— 1 — — 2 12— 2 1—— 1 2 2 25 40 32 40 10 41 18 27 16 8 25 18 53 54 Se We a ee Sd IRA ERS, 2.3.3. NRN eet wasmot 4. 3 a ies ee — (ee aloes A ee ee E ee | 49 71 62 77 31 87 48 71 82 41 54 27 98 86 teils sind es mediterrane Arten (69), und die meisten von ihnen gehòren zur Untergattung Tipula (Lunatipula) (66). Die Balkanhalbinsel und das italienische Festland Die Balkanhalbinsel und Italien haben viele Arten gemeinsam; es sind dies aber fast aus- nahmslos Arten mit Verbreitung bis in die euro- paischen Tiefebenen und kaum Arten mit Ver- breitung in den Gebirgen oder in den mediterra- nen Gebieten (Theowald & Oosterbroek, 1983, 1984). In den beiden letzten Kaltzeiten muf es fur die Arten der Tiefebenen ein gemeinsames italo-balkanisches Refugium gegeben haben (Theowald & Oosterbroek, 1984). Verbreitung und Verwandschaftsbeziehungen deuten aber daraufhin, daß nach der letzten Kaltzeit kaum noch Arten von Italien nach der Balkanhalbinsel oder umgekehrt gekommen sind. Die Balkanhalbinsel und Kleinasien Die Balkanhalbinsel hat nur wenige Arten mit dem naheliegenden artenreichen Kleinasien ge- meinsam. Von den Arten der europaischen Tief- ebenen finden sich 16 nicht nur auf der Balkan- halbinsel sondern auch in Kleinasien, oder sie THEOWALD & OOSTERBROEK: Westpalaearktischen Tipuliden. VII 9 haben dort eine nachstverwandte Schwesterart (Theowald & Oosterbroek, 1983). Von den Ge- birgsarten hat nur eine in Kleinasien ein Schwesterart (Theowald & Oosterbroek, 1985). Von den mediterranen Arten haben sich rezent neun von Kleinasien bis nach der Balkanhalbin- sel verbreitet (Tabelle 4B, 3b) und drei von der Balkanhalbinsel nach Kleinasien (bimacula, soo- si und italica errans). Insgesamt kennt man von der Balkanhalbinsel und von Kleinasien zusam- men etwa 300 Arten, von denen nur 29 in bei- den Gebieten vorkommen. Von der Balkanhalb- insel und Italien hingegen kennen wir insgesamt etwa 250 Arten, von denen fast 100 in beiden Gebieten vorkommen. 2. Die Verbreitung der Tipuliden ım balkanischen Raum Aus den Tabellen 1—4 geht hervor, daß es auf der Balkanhalbinsel einen Unterschied gibt zwischen der Verbreitung der mediterranen und jener der nicht-mediterranen Arten. Letztge- nannte sind ziemlich gleichmäßig über das gan- ze Gebiet verbreitet, die mediterranen Arten dagegen finden sich vorwiegend in der West- hälfte der Balkanhalbinsel. Eine Auszählung von Tabellen 1—3 ergibt: Westbalkan (Jugoslawien, Albanien, Grie- chenland) mediterrane Arten: 85 der insgesamt 88, d.h. 97% nicht-mediterrane Arten: 91 der insgesamt 113, d.h. 81% Ostbalkan (Rumänien, Bulgarien) mediterrane Arten: 16 der insgesamt 88, d.h. 18% nicht-mediterrane Arten: 102 der insgesamt 113, d.h. 90% Der Westbalkan ist somit am reichsten an Ti- puliden. Er hat nicht nur viele nicht-mediterra- ne Arten, die er mit dem Ostbalkan gemeinsam hat, sondern auch viele mediterrane Arten, von denen nur wenige auch im Ostbalkan vorkom- men. Es gibt aber auch Unterschiede zwischen Nord- und Südbalkan. In Tabelle 5 (vgl. auch Karte 2) wird die Balkanhalbinsel in fünf größe- re Gebiete unterteilt: Nordwest-, Mittelwest- und Südwestbalkan und Nordost- und Südost- balkan. Für jedes dieser Teilgebiete ist die Fau- nenzusammensetzung absolut und prozentuell verzeichnet. Im Westbalkan finden sich vom Norden nach dem Süden absolut und prozen- tuell immer mehr mediterrane und immer weni- ger nicht-mediterrane Arten. Im Nordwestbal- kan ist jedoch noch ein Drittel der Arten medi- terran und im Südwestbalkan ein Drittel der Arten nicht-mediterran. Im Ostbalkan sind pro- zentuell die Unterschiede zwischen beiden Ge- bieten viel kleiner. Beide Faunen sind vorwie- gend nicht-mediterran. Die Zahl der neun medi- terranen Arten im Nordostbalkan (Tabelle 5) ist aber noch flattiert, denn von diesen neun Arten, kommen vier nur im Banater Gebirge an der ju- goslawisch-rumänischen Grenze vor (Tabelle 3 mit . statt +) und sind nur fünf Arten recht ost- balkanisch. Der ostbalkan, ganz besonders aber der Nordostbalkan, schließen hinsichtlich Fau- nenzusammensetzung eng an das mitteleuropäi- sche Gebiet an, der Westbalkan dagegen wird nach dem Süden hin immer deutlicher mediter- ran. 3. Das balkanische Zentrum mediterraner Lunatipula-Arten Die Untergattung Lunatipula ist in ihrer Ver- breitung hauptsächlich auf die mediterranen Gebiete der Holarktis beschränkt. Im mediter- Tabelle 5. Faunenzusammensetzung der fünf Teilgebiete der Balkanhalbinsel. Nordwest- Mittelwest- Südwest- Südost- Nordost- balkan balkan balkan balkan balkan mediterrane Arten absolut: 87 33 46 61 12 9 prozentuell: 43% 28% 41% 66% 18% 9% nicht-mediterrane Arten (Arten der Tiefebenen) absolut: 72 62 53 DD 40 63 prozentuell: 36% 53 % 47% 23% 61% 60% (Arten der Gebirgen) absolut: 42 22 13 10 14 52 prozentuell: 21% 19% 12% 11% 21% 31% Total: 201 Arten 117 112 93 66 104 10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 1, 1986 one 1 A “> Kroatien a, ry FE x 1 Noe Banane > ==. € à È SH Bosnien À & i‘ Q Oy \ i by “Herzego- > I \ wina 5 Q LES DR fy Qu re Seroen SE td s Montenegro en 77 Karte 1. Balkanhalbinsel mit Teilgebieten. ranen Raum der Palaearktis finden sich 250 Ar- ten (Theischinger & Theowald, 1981). Verhalt- nısmäßig wenige dieser 250 Arten finden sich im iberischen und im italienischen Gebiet (Theowald & Oosterbroek, 1980, 1981, 1984). Die meisten sind auf der Balkanhalbinsel oder ın Kleinasien verbreitet. Unter den mediterranen Arten der Balkan- halbinsel sind 79 Lunatipula-Arten, von denen neun rezent eingewandert sind und 70 zur ursprünglichen Fauna gehören (Tabelle 3). Letztgenannte finden sich alle im Westbalkan, einige überdies im Ostbalkan (leandros ist nur bekannt vom Banater Gebirge an der Grenze von Rumänien und Jugoslawien und wird zur Fauna des Westbalkan gezählt). Im Westbalkan Na “__---77 Griechenland ed a ; ZU ---- NI Dobrudscha kommen die meisten Arten in Griechenland vor, und zum Teil reichen sie von dort mehr oder weniger weit nach Jugoslawien. Viel weni- ger Arten sind in ihrer Verbreitung auf Jugosla- wien beschrankt oder machen den Eindruck, daß sie sich von dort mehr oder weniger weit nach Griechenland verbreitet haben (Tabelle 3). Von den 70 zur ursprünglichen Fauna gehöri- gen Arten gehören 40 (57%) zu drei Artengrup- pen mit fast rein balkanischer Verbreitung: die clio-, truncata- und fascingulata-Gruppe. Insge- samt kommen von diesen drei Artengruppen nur drei Arten im kleinasiatischen Raum vor. Es ist wohl sicher, daß diese drei Artengruppen sich auf der Balkanhalbinsel differenziert haben, und daß dreimal eine Art von dort bis nach THEOWALD & OOSTERBROEK: Westpalaearktischen Tipuliden. VII 11 Nordostbalkan Sudostbalkan Sudwest- balkan N A W Karte 2. Hauptgebiete der Balkanhalbinsel mit Faunenzusammensetzung (schwarz: Gebirgsarten; schraffiert: Arten der Tiefebenen; weiß: mediterrane Arten). Kleinasien gelangt ist. Insgesamt 25 der 70 Ar- ten (36%) gehören zu sechs Artengruppen mit vorwiegend kleinasiatischer Verbreitung: acu- minata-, macroselene-, livida-, peliostigma-, brunneinervis-, und lunata-Gruppe. Im klein- asiatischen Raum gibt es von diesen Gruppen noch etwa 85 weitere Arten. Die Verwandt- schaftsbeziehungen sofern bekannt, zwischen den Arten der genannten Artengruppen, lassen den Schluß zu, daß im Laufe der Zeit mehrmals Arten von Kleinasien auf die Balkanhalbinsel gekommen sind, sich dort zu endemischen Ar- ten differenziert haben und sich zum Teil dort wieder weiter in Arten aufgespalten haben. Ins- gesamt nur fünf Arten gehören zu drei Arten- gruppen mit Hauptverbreitung im westmediter- ranen Gebiet, auf Kreta oder im Kaukasus (Ta- belle 3). Obwohl sowohl der Westbalkan, insbesonde- re Griechenland, als auch Kleinasien ihre ende- mische Lunatipula-Fauna haben, hat es zwi- schen den beiden Gebieten dann und wann Austausch gegeben. Der kleinasiatische Einfluß auf der Balkanhalbinsel ist aber deutlich viel größer als jener der Balkanhalbinsel in Klein- asıen. Dasselbe finden wir auch bei den rezent ein- und ausgewanderten Arten, die noch in bei- den Gebieten unverändert vorkommen: sieben Lunatipula-Arten haben sich rezent von Klein- asıen bis auf die Balkanhalbinsel verbreitet (Ta- 12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 1, 1986 belle 3 mit K), dagegen nur zwei von der Bal- kanhalbinsel bis nach Kleinasien (bimacula und 50051). Wie die Iberische Halbinsel und Nordafrika, liegen auch die Balkanhalbinsel und Kleinasien geographisch nahe zusammen, und doch hat sich in beiden Fallen in jedem dieser Gebiete eine endemische Lunatipula-Fauna entwickelt, obwohl es in beiden Fallen auch Auswechslung gegeben hat. Die Lunatipula-Fauna der beiden ostmediterranen Gebiete ist jedoch viel reicher als die der beiden westmediterranen Gebiete. Im ostmediterranen Gebiet gibt es zwei Zentren für Artbildung mediterraner Lunatipula-Arten: der Westbalkan, insbesondere Griechenland, und Kleinasien. ZUSAMMENFASSUNG Die Verbreitung der 201 Tipuliden-Arten, die auf der Balkanhalbinsel nachgewiesen sind, wird beschrieben und analysiert. Folgende Schlüsse werden gezogen. Die meisten Arten der mittel- und westeuro- päischen Tiefebenen und Gebirge kommen auch ziemlich gleichmäßig verbreitet über die ganze Balkanhalbinsel vor; in Mittel- und Südgrie- chenland gibt es aber etwas weniger Arten als weiter im Norden. In Jugoslawien und zumal in Griechenland finden sich überdies auch viele mediterrane Ar- ten, die im Ostbalkan fast vollständig fehlen. Die Tipuliden-Fauna des ostbalkan ist somit deutlich mittel- und westeuropäisch, die des Westbalkan dagegen hat stärker mediterranen Charakter. Das italienische Festland und die Balkanhalb- insel haben viele Arten der europäischen Tief- ebenen gemeinsam, nicht aber Arten der Gebir- ge und mediterrane Arten. Für die Arten der europäischen Tiefebenen hat es im Eiszeitalter wohl ein italo-balkanisches Refugium gegeben. Die Balkanhalbinsel und Kleinasien haben kaum Arten gemeinsam. Zwischen beiden Ge- bieten hat es aber im Laufe der Zeit wohl Aus- tausch gegeben, im Zuge dessen insbesondere Lunatipula-Arten sich von Kleinasien bis auf die Balkanhalbinsel verbreitet haben aber kaum umgekehrt. Der Westbalkan und Kleinasien sind im me- diterranen Gebiet der Palaearktis wohl die be- deutendsten Gebiete für Artbildung mediterra- ner Lunatipula-Arten gewesen. SUMMARY The distribution of the 201 tipulid species from the Balkan peninsula is presented and ana- lyzed, with the following conclusion. Most lowland or mountain species of central and western Europe are distributed throughout the Balkan peninsula but with a lesser amount of species in central and southern Greece than further to the north. Jugoslavia and Greece count many mediterra- nean species. This element is virtually lacking in the east Balkan. This part of the area is much more central and western european, whereas the west Balkan has a stronger mediterranean cha- racter. The Italian mainland and the Balkan peninsu- la have many European lowland species in com- mon. This is not the case for the mountainous and mediterranean species. An Italo-Balkanian refugium is postulated for the lowland species during the latest glaciations. The Balkan peninsula and Asia Minor hardly : have species in common. Exchange during cer- tain periods must have occurred and especially within Lunatipula, species of which migrated from Asia Minor to the Balkan peninsula and, to a far lesser degree, in the opposite direction. The west Balkan and Asia Minor are of the Mediterranean part of the Palaearctic the two important regions for speciation within Lunati- pula. LITERATURLISTE Erhan, E., 1959. Contributii la cunoasterea faunei de tipulide (Diptera-Tipulidae) din Masivul Bucegi si cursul superior al riului Prahova. — Studii Cerc. Biol., ser. Biol. Anim. 11: 217—225. Erhan, E., 1962a. Date noi cu privire la fauna de tipu- line (Diptera-Tipulidae) din R.P.R. — Studii Cere. Biol., ser. Biol. Anim. 14: 91—109. Erhan, E., 1962b. Contributu la studiul faunei de Ti- pulinae (Diptera-Tipulidae) din R.P.R. — Studii Cerc. Biol., ser. Biol. Anim. 14: 351—370. Erhan, E., 1971. Noi specii de Tipulinae (Diptera-Ti- pulidae) din fauna Romaniei. — Studii Cerc. Biol., ser. Zool. 23: 537—550. Erhan, E., 1973. Noi contributii la cunoasterea tipuli- nelor (Diptera-Tipulidae) din Romania. — Studi Cerc. Biol., ser. Zool. 25: 437—445. Erhan, E., 1976. Contributii la studiul tipulinelor din subgenul Lunatipula (Diptera-Tipulidae) din Ro- mania. — Studii Cerc. Biol., ser. Biol. Anim. 28: 13—20. Erhan, E., 1959. Die Entwicklungsstadien von Tipula soosi und T. peliostigma. — Tijdschr. Ent. 102: 217—221. Erhan, E., & Br. Theowald, 1961. Tipulidae of Rou- mania (Dipt., Nematocera). — Ent. Ber. Amster- dam 21: 245—252. THEOWALD & OOSTERBROEK: Westpalaearktischen Tipuliden. VII 13 Kowarz, P., 1873. Beitrag zur Dipteren-Fauna Un- garns. — Verh. zool. bot. Ges. Wien 23: 453— 464. Mannheims, B., 1954. Die Tipuliden Griechenlands (Dipt.). — Bonn. zool. Beitr., Sonderband 1954, I: 149—182. Mannheims, B., 1966. Ergebnisse der Albanien-Expe- dition 1961 des Deutschen Entomologischen In- stitutes. 56. Beitrg. Diptera: Tipulidae. — Beitr. Ent. 16: 489—502. Mannheims, B., & Br. Theowald, 1951—1980. Tipuli- dae. — Fliegen palaearkt. Reg. III, 5, 1. — Schweizerbart, Stuttgart. Savtshenko, E. N., 1961—1983. Tipulidae. — Fauna SSSR, Insecta, Diptera II, 1—2 (N.S.127, 1983), IL, 3 (N.S.79, 1961), II, 4 (N.S.89, 1964), II, 5 (N.S.105, 1973). — Nauka, Moskou, Leningrad. Simova, D., 1959. Beitrag zur Kenntnis der Tipuliden und Limnobiiden von Mazedonien. — Fragm. Balc. 2: 125—135. Simova, D., 1960. Drugi prilog poznavanju tipulida 1 limnobiida makedonije. — Fragm. Balc. 3: 51— 63. Simova, D., 1962. Tipulidae nekih krajeva jugoslavije. — Fragm. Balc. 4: 101—105. Simova, D., 1964. Neke vrste familije Tipulidae nove za jugoslaviju. — Fragm. Balc. 5: 49—54. Simova, D., 1972. Due nove vrste iz familije Tipulidae (Diptera). — Zast. Bilja 23:333—339. . Simova, D., 1973. Contribution to the study of Crane | | | Flies (Tipulidae, Diptera) in Yugoslavia. —Fragm. Balc. 9: 149—153. Simova, D., 1974. Prilog poznavanju faune Tipulidae (Diptera, Nematocera) jugoslavije. — Zast. Bilja 25; 25—30. Simova, D., 1976. Prilog proucavanju faune Tipulidae (Diptera) sr srbije. — Arh. biol. nauka. Beograd 28: 147—152. Simova, D., 1978. Dolichopeza graeca Mnhs. (Dipte- ra, Tipulidae) nov vid sa faunata na jogoslavija. — Fragm. Balc. 10: 135—138. Simova, D., & I. Sivec, 1978. Prispevek k poznavanju faune koseninarjev (Diptera, Tipulidae) v slove- niji. — Biol. vestn. Ljubljana 26: 183—186. Simova, D. & B. Todorovski, 1977. Tipulidae (Dipte- ra) ulovljene na svetlostne klopke. — Tutun To- bacco 5—6: 249—260. Simova, D. & M. Vukovic, 1981. The results of Crane Flies (Diptera, Tipulidae) studies in Yugoslavia. — Acta ent. Jugoslavica 17: 13—119. Simova, D. & Vukovic, 1983. Usko rasprostranjene vrste faune Tipulidae (Diptera) jugoslavije. — Drugi simpozijum o fauni SR Srbije, Zbornik: 99—102. — Beograd. Simova, D. & Vukovic, 1983. Znacaj muskog genital- nog aparata za determinacju vresta familije Tipuli- dae (Diptera). — Acta ent. Jugoslavica 19: 27—32. Strobl, G., 1897. Siebenbürgische Zweifligler. — Verh. Mitt. siebenb. Ver. Naturwiss., Hermann- stadt 46: 11—48. Strobl, G., 1900. Dipterenfauna von Bosnien, Herze- govina und Dalmatien. — Wiss. Mitt. Bosnien Herzegovina 7: 552—670. Strobl, G., 1904. Neue Beitrage zur Dipterenfauna der Balkanhalbinsel. — Wiss. Mitt. Bosnien Her- zegovina 9: 519 - 581. Szilady, Z., 1934. Beitrage zur Dipterenfauna Bulga- riens. — Izv. bulg. ent. Druzh. 8: 145—151. Thalhammer, J., 1900. Ordo Diptera in Fauna Regni Hungariae III, Arthropoda. Ed. sep. 1—76. Theischinger, G., 1977. Neue Taxa von Lunatipula Edwards aus der mediterranen Subregion der Pa- laarktis (Diptera, Tipulidae, Tipula Linnaeus). — Beaufortia 26: 1—38. Theischinger, G., 1979a. idem, 1. Beaufortia 28: 121—150. Theischinger, G., 1979b. idem, 2. Fortsetzung. — Beaufortia 29: 275—308. Theischinger, G., 1980. idem, 3. Fortsetzung. —Beau- fortia 30: 17—29. Theischinger, G., 1982. idem, 4. Fortsetzung. — Beaufortia 32: 79—96. Theischinger, G., 1983. Uber eine Aufsammlung von Savtshenkia Alexander, 1965, aus dem östlichen Mittelmeerraum (Diptera: Tipulidae). — Ent. Ber. Amsterdam 43: 26—28. Theischinger, G. & Br. Theowald, 1981. Katalog der Untergattung Lunatipula Edwards, 1931, s.l. — Versl. techn. Geg. Inst. Tax. Zool. Univ. Amster- dam 27: 1—55. Theowald, Br., & P. Oosterbroek, 1980. Zur Zoogeo- graphie der westpalaearktischen Tipuliden. I. Die Tipuliden von Nordafrika. — Beaufortia 30: 179—192. Theowald, Br., & P. Oosterbroek, 1981. idem, II. Die Tipuliden der Iberischen Halbinsel. — Beaufortia 31:31—50. Theowald, Br. & P. Oosterbroek, 1983. idem, III. Die Tipuliden der europaischen Tiefebenen. — Bonn. zool. Beitr. 34: 371— 394. Theowald, Br., C. Dufour & P. Oosterbroek, 1982. idem, IV. The Tipulidae of Corsica and Sardinia with a note on Dolichopeza fuscipes Bergroth. — Mitt. schweiz. ent. Ges. 55: 317—332. Theowald, Br., & P. Oosterbroek, 1984. idem, V. Die italienischen Tipuliden. — Fragm. Entomol., Ro- ma 17: 245—291. Theowald, Br., & P. Oosterbroek, 1985. idem, VI. Die Tipuliden der montanen, alpinen und borealen Gebiete. — Bonn. zool. Beitr. 36: 185—220. Fortsetzung. — : hints ta N erie N ia a Ay MES te RN: Be di ITS La Ur ys Pt Vor [fo a) Lg Dh ee U DEEL 129 AFLEVERING 2 1986 cs TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR # | 4 74 DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING & 7 Z INHOUD Tuomas PAPE. — A phylogenetic analysis of the Woodlouse-flies (Diptera, Rhino- phoridae), pp. 15—34, figs. 1—30. Tijdschrift voor Entomologie, deel 129, afl. 2 Gepubliceerd 10-XI-1986 A PHYLOGENETIC ANALYSIS OF THE WOODLOUSE- FLIES (DIPTERA, RHINOPHORIDAE) / by THOMAS PAPE Zoological Museum Copenhagen, Denmark ABSTRACT The Rhinophoridae are redefined on the basis of the apomorphic structure of the aedea- gus. Evidence is provided for the exclusion of four genera, viz., Angioneura Brauer & Ber- genstamm, 1893, Melanomya Rondani, 1856, Morinia Robineau-Desvoidy, 1830, and Ter- mitoloemus Baranov, 1936; all four are transferred to the Calliphoridae. The genera of Rhi- nophoridae are analyzed phylogenetically with the aid of the results of the present investigation and the sparse information available on the morphology of the larval stages. The structure of the aedeagus provides several set-defining characters and the aedeagus of many species is depicted for the first time. Czrillia Rondani, 1856, is proposed as a synonym of Phyto Robineau-Desvoidy, 1830. INTRODUCTION Within the calyptrate flies the species with a | row of bristles on the meron (hypopleuron) | constitute a well-corroborated monophyletic | group, | Girschner (1893), Calliphoroidea sensu Hennig | (1958), Tachinidae (sensu lato) sensu Griffiths | (1972), or Oestroidea sensu McAlpine et al. | (1981)). Most recent authors, including the pre- | sent, accept five major groups in the Tachinoi- | dea, viz., Oestridae (sensu lato), Calliphoridae, | Sarcophagidae, Tachinidae, and Rhinophoridae. the Tachinoidea (Tachinidae sensu Although Crosskey (1965) restricts the name | Tachinoidea to the Calliphoridae, Sarcophagi- | dae, Tachinidae, and Rhinophoridae, synapo- | morphies not shared by the Oestridae (which i would be their sister group), to my knowledge | have not been provided for these four families by any author. The family Stackelbergomyiidae Rohdendorf, 1948, was obviously established because no evi- | dence for incorporating the single aberrant spe- | cies into any of the existing families could be | found. An investigation by Herting (1981) sug- | gests that it should be included in the Tachini- || dae. More interesting are the Neotropical Me- || sembrinellinae (Calliphoridae). Crosskey (1965) \ is of the opinion that an improved classification \ of the Tachinoidea (in his definition as given above) would result if “peculiar groups such as O ee were treated as families” (p. 43). Guimaraes (1977) follows this recommen- dation and raises the group to family status: Mesembrinellidae, founding his decision on five “consistent differences” between Mesembrinel- linae and the remaining Calliphoridae. These differences corroborate the monophyly of the Mesembrinellinae, but the Calliphoridae sensu Guimaraes are characterized solely on symple- siomorphies and fail to support a family status of the Mesembrinellinae. An argument for split- ting'up the Calliphoridae would be that the sim- ple, non-opercular lappet of the mesembrinel- line metathoracic spiracle is plesiomorphic, as this would separate the Mesembrinellinae (still monophyletic) not from the Calliphoridae but from all other Tachinoidea, the monophyly of which would be corroborated by their opercu- lar metathoracic spiracle. This may be the rea- son for Crosskey’s (1965: 43) note that the Me- sembrinellinae “may not be Tachinoidea at all”. I hesitate to place the Mesembrinellinae as sister group to all other Tachinoidea and prefer to treat them as Calliphoridae. The structure of the mesembrinelline aedeagus with strong, for- wardly curved dorsolateral processes (paraphal- li) seems a reasonable synapomorphy with the Calliphoridae (and perhaps with the Rhinopho- ridae?). A small digression may be made here, brought about by the recent (and past) dis- agreement of family status criteria. Some au- thors, e.g., Steyskal (1974) and Hackman & Vaisanen (1982), have mentioned the inconsis- tency of Griffiths’ (1972) splitting of the Musci- dae sensu Hennig (1958, 1965) into Muscidae and Fanniidae when he unites all tachinoid flies in a single family: Tachinidae (sensu lato). 16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 However, although Hennig (1965) states that: “Eine der sichersten Feststellungen, vielleicht die gesichertste, die man über das phylogene- tische System der Muscidae treffen kann, ist die, dass zwischen den Fanniinae auf der einen seite und der Gesamtheit aller übrıgen Muscidae... ein Schwestergruppenverhältnis besteht” (p. 9), he does not bring conclusive evidence of the monophyly of the “Muscidae sensu lato”. Therefore, a separation is to be preferred. If the tachinoids are considered a monophyletic group, they are best treated in common when used for outgroup comparison, and the formal rank — whether family or superfamily — is of minor importance in a phylogenetic sense. Only the ranking of the group relative to the other Calyptratae is important as this constitutes a phylogenetic hypothesis. The monophyly of the Tachinoidea seems fairly corroborated. Griffiths (1972) mentions the following synapomorphies with respect to the groundplan of the Calyptratae: (1) hypopleuron with strong bristles below metathoracic spiracle, (2) eighth sternum (©) entire, (3) vein m,,, sharply bent towards r,,, apical- ly, (4) anal vein not reaching wing margin, (5) sixth tergum (d) shortened, less than half as long as 5th tergum, (6) eighth tergum vestige (2) lost. The loss of the “eighth tergum vestige” in males is based on a questionable interpretation of a median ventral sclerotization in the postab- domen of some Anthomyiidae and Scatophagi- dae (Griffiths, 1972: fig. 61); this sclerotization more likely is a secondary acquisition. Another character which may be autapomor- phic to the Tachinoidea is: (7) lappet of metathoracic spiracle divided, pos- terior lappet shaped as an operculum. This opercular metathoracic spiracle, absent in all other calyptrates, is present in the majority of the Tachinoidea; the non-opercular metatho- racic spiracle present in the Mesembrinellinae and a few other Calliphoridae, some groups of Tachinidae, Macronychiinae of the Sarcophagi- dae, almost all Rhinophoridae, and many Oes- tridae (sensu lato) may be secondarily derived. The sister group relations of the Tachinoidea within the Calyptratae are still largely unsolved, and the characters mentioned by Griffiths are not necessarily autapomorphies for the Tachi- noidea, viz., items 4 and 5 mentioned above, which also occur among other calyptrate groups. A shortened anal vein (A,) is character- istic of both the Muscidae and Fanniidae. A few genera in the Tachinoidea (e.g., some Oestridae (sensu lato) and Tachinidae, Bengalia Robi- neau-Desvoidy in the Calliphoridae) possess an extended anal vein, a character which both Hennig (1958) and Griffiths (1972) consider to be secondary. It is interesting, however, that Andersen (1982) reports aerial swarming of male Siphona Meigen as the first example within the Tachinidae and suggests (Andersen, 1982, 1983) that an extended anal vein may be assig- nable to the groundplan of the Tachinidae (and then possibly to all the Tachinoidea). The shortened abdominal tergum 6 in males is of general occurrence in the Muscidae and An- thomyiidae as well. The Rhinophoridae are typical members of the Tachinoidea as defined above (fig. 1), but. the affinities to other tachinoid families are still unclear. Many earlier authors placed the rhino- phorids with the blow-flies and flesh-flies in a Calliphoridae (sensu lato), but in a phylogenetic sense this constitutes an entirely unacceptable non-group arising by the splitting off of the flies possessing a swollen subscutellum — the Tachi- nidae. Mesnil (1939) derived most of the subfa- milies of Tachinidae from different rhinophorid stocks, thereby rendering the Rhinophoridae paraphyletic (and the Tachinidae polyphyletic), but at present most authors give the Rhinopho- ridae family rank, acknowledging their uniqueness and the present lack of evidence for a closer relation to any of the other tachinoid families. Kugler (1978), and especially Crosskey Fig. 1. Stevenia deceptoria (Loew); a typical wood- louse-fly. Pare: Rhinophoridae 17 (1977), give a more detailed review of previous differences of opinion regarding the family affi- nities. LARVAL BIOLOGY AND MORPHOLOGY Although the family is small, an unambiguous demarcation of the Rhinophoridae has not been possible. This is due in part to the existence of deviating tropical forms, e.g., Bequaertiana Curran, and in part to an external morphology intermediate between that of typical callipho- rids and typical tachinids. More important, however, is the lack of information concerning the morphology and biology of the larval stages. All known first-stage larvae possess a distinctive cephalopharyngeal skeleton with the anterior part of the pharyngeal sclerite greatly elongated and with two or more teeth on the dorsal arc of the mandibles — evidently synapomorphic characters. The larval habit of parasitizing woodlice (Isopoda) is likewise unique to the Rhinophoridae, and interesting insofar as very few biological relationships between Diptera and Crustacea are known (see Roubaud, 1903; Mercier, 1921; Oldroyd, 1964, and Burger et al., 1980). Only seven genera of rhinophorids actually have been recorded as woodlouse parasites, viz., Stevenia Robineau-Desvoidy, Tricogena Ron- dani, Rhinophora Robineau-Desvoidy, Melano- phora Meigen, Paykullia Robineau-Desvoidy, Phyto Robineau-Desvoidy, and Cirillia Ronda- ni (note that Cirillia is a synonym of Phyto, see discussion below). Specific host records for the Palaearctic species are given by Herting (1961) with supplements in Kugler (1978). Parker (1953) mentions breeding of the introduced Melanophora roralis (Linnaeus) in Brazil. No host records exist for any of the Nearctic, Afro- tropical, or Oriental species. Table 1. List of non-isopod hosts of the Rhinophoridae. There has been some doubt as to whether the Rhinophoridae could be parasites in inverte- brates other than isopods, and the tendency has been to disregard any such record. Obviously, the report of Melanophora helicivora Goureaux being bred from the gastropod Helicella con- spurcata (Draparnaud) is based on a mis-identi- fication. As judged from the description and drawings (Goureaux, 1843: figs. 1, 2), the spe- cies does not belong to Melanophora at all, but may be a calliphorid. Lundbeck (1927) mentions a specimen of Melanophora roralis bred from egg-cocoons of the spider Araneus cornutus Clerck. I have seen this specimen, a female deposited in the Zoolog- ical Museum, Copenhagen, and it is correctly identified by Lundbeck. In addition to this there are several reports of rhinophorids parasitizing insects (table 1), and it is probable that rhinophorids occasionally (acci- dentally?) may parasitize arthropods other than isopods. Very little has been written on the morpholo- gy of the larvae of the woodlouse-flies. Thomp- son (1934) treated in detail the larval stages of eight species, viz., Paykullia maculata (Fallén), Phyto angustifrons (Rondani), Phyto discrepans (Pandellé), Phyto melanocephala (Meigen), Melanophora roralis, Stevenia atramentaria (Meigen) (as species B), Tricogena rubricosa (Meigen), and Rhinophora lepida (as species A). However, Thompson obtained all his material from dissections of woodlice as most of his at- tempts to obtain eggs from female flies caught in the wild and hatch these to first-instar larvae failed. Furthermore, he often assumed that rhi- nophorid larvae from a single colony of wood- lice were conspecific. This has resulted in some erroneous identifications in his earlier works (Thompson, 1917, 1920; corrected in 1934: parasite/predator specimens host reference Melanophora roralis 19 eggs of Araneus cornutus Clerck Lundbeck (1927) (Araneae) ? ?Pyralıs farinalis (Linnaeus) Bezzi & Stein (1907) (Lepidoptera, Pyralidae) Stevenia umbratica ? Callidium violaceum Linnaeus Bezzi & Stein (1907) (Coleoptera, Cerambycidae) Rhinophora lepida 18 Paranthrene tabaniformis (Rottemburg) Kolubajıv (1962) (Lepidoptera, Aegeriidae) 1d Saperda carcharias (Linnaeus) Kolubajıv (1962) (Coleoptera, Cerambycidae) Neh Ne Rhinomorinia sarcophagina Malacosoma neustria (Linnaeus) Kolubajıv (1962) (Lepidoptera, Lasiocampidae) 18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 380). Of the first-instar larva of Phyto angustif- rons Thompson had only a single defective specimen (the cephalopharyngeal skeleton and a skin fragment). The depicted cephalopharyngeal skeleton (Thompson, 1934: pl. 19, fig. 47) is of the heavy, sclerotized type found in Stevenia, Tricogena, and Rhinophora and very unlike the cephalopharyngeal skeleton of Paykullia, Mela- nophora, and other species of Phyto. The first-stage larva assigned to Phyto angus- tifrons probably belongs to another species (very likely a Stevenia). The cephalopharyngeal skeleton of the second- and third-stage larva of P. angustifrons (Thompson, 1934: pl. 20, figs. 48, 56) is more in accordance with that of Phyto species. Bedding (1973), in an extract of his Ph. D. thesis, described eggs and larval stages of all English species — actually the same species as those described by Thompson (1934) except for P. angustifrons. The larvae, especially first instars, possess several features which are very useful in a phylogenetic context, but at present the larval stages are known for only a small fraction of the species described. In addition, the uniqueness of many of the features makes any outgroup comparison almost inapplicable in the distinction between apomorphic versus ple- siomorphic larval characters within the family. The first-stage larvae known at present com- prise two distinct groups (see figs. 8—44 in Bedding, 1973): A. Phyto, Paykullia, Melanophora (1) mandibles with normal degree of sclerotiza- tion, with three or more small teeth on the dorsal arc, (2) elongated anterior part of pharyngeal scle- rite with an incision, (3) setal bases unmodified, (4) posterior end of larva highly modified for supporting the larva in erect posture; with a dorsal tongue, terminal sac-like lobes, and ventral ridges. B. Stevenia, Tricogena, Rhinophora (1) mandibles heavily sclerotized, with two strongly developed teeth, (2) elongated anterior part of pharyngeal scle- rite without an incision, (3) setal bases protruded into proleg-like struc- tures, (4) posterior end of larva simple, with inflated ventral vesicles. Bedding notes that the two morphologically distinct groups of first-stage larvae possess dif- ferences in their biology (referring to a paper (in prep.) which unfortunately has not yet been published). The toothed mandibles of the first-stage lar- vae are probably an adaptation for penetrating the body wall of the host, analogous to the ser- rate median tooth of tachinid larvae which enter the host through a strongly sclerotized cuticle (Clausen, 1940: fig. 210 A). This character is clearly an autapomorphy for the Rhinophoridae as toothed mandibles occur very sporadically in other Tachinoidea, e.g., the warblefly of the lechwe antelope (Howard, 1980). The two types of cephalopharyngeal skeleton can not be separated into an apomorphic and a plesiomorphic state at present; indeed, it is pos- sible that both types are apomorphic with re- spect to the groundplan of the Rhinophoridae, but this may be the least parsimonious solution to the problem. The proleg-like setal bases must be consid- ered an apomorphic character as these are ab- sent in the majority of the Tachinoidea and nothing indicates their suppression in other rhi- nophorids. This character is found in Stevenza, Tricogena, and Rhinophora and may be a syna- pomorphy of the Stevenia group (see discussion below), thereby corroborating the monophyly of this group. The two types of modified posterior end of the first-stage larva present a problem some- what analogous to that of the cephalopharyn- geal skeleton. However, until more information on the sister group relations of the Rhinophori- dae within the Tachinoidea becomes available, it is reasonable to assume that the sister group possesses first-stage larvae with unmodified posterior ends. The terminal lobes, the dorsal tongue, and the free, posteriorly oriented ven- tral ridges will then be apomorphic characters, and the terminal lobes will be the apomorphic homologues of the inflated vesicles. This will corroborate the hypothesis that Paykullia, Phy- to, and Melanophora are part of a monophyletic group (the Phyto group) not containing Steve- nia, Tricogena, or Rhinophora. RECOGNITION OF THE RHINOPHORIDAE Crosskey (1977: 7) gives an excellent dis- cussion of the status and recognition of the fam- ily, but he admits that his recognition couplet does not ensure a certain identification. A fur- ther complication is the recently described ge- nus Baniassa Kugler. This genus has a well-de- veloped metathoracic opercular spiracle, but the absence of a distinct operculum has hitherto Pare: Rhinophoridae 19 provided one of the most important single char- acters for rhinophorid recognition. Besides the structure of the metathoracic spiracle, the char- acters most helpful in recognizing the family have been the tongue-shaped or oval lower ca- lypteres which are widely removed from the scutellum, the bend of vein M which never is greatly concave, and the combination of bare prosternum, proepisterna, greater ampullae, postalar walls, laterotergites, and supra-squamal ridges. In the majority of the Tachinoidea the struc- ture of the aedeagus (and other structures of the terminalia) provides important characters in the diagnostic segregation of species and is often used in the construction of evolutionary trees and in the definition of taxonomic categories above the species level. Some illustrative exam- ples are the works of Mueller (1926) on the Ta- chinoidea, Roback (1954) on the Sarcophaginae, Verbeke (1962) on the Tachinidae, Kurahashi (1966) on the Luciliinae, Lehrer (1970) on the Calliphoridae, and Lehrer (1973) on Sarcophaga (sensu stricto). The distiphallus of male rhino- phorids, however, is seldom depicted, not even in the revisions of the Palaearctic (Herting, 1961) and Afrotropical (Crosskey, 1977) spe- cies, and the information stored in this structure is largely unknown. Mueller (1926) made an early attempt to construct a “Stammbaum... auf Grund der Penisform” of the Tachinoidea, but only a few rhinophorids were included and the drawings are more or less incorrect. Séguy (1941) made a preliminary division of the Rhi- nophoridae (as a subfamily of the Calliphoridae, sensu lato) into four groups on the basis of the male genitalia, but he dissected only a few rep- resentatives and his definition of the (sub)family included several tachinid, sarcophagid, and cal- liphorid genera. The structure of the aedeagus may provide additional characters to be used in the recogni- tion of the family; and in order to use this struc- ture in a redefinition of the family and in the re- construction of the phylogeny at the generic level, the following hypothetical groundplan of the tachinoid aedeagus is accepted (terminology as in Hennig, 1976 and McAlpine et al., 1981) (fig. 2). Like most other calyptrate flies a well-devel- oped basiphallus, distiphallus and epiphallus are present. The distiphallus is more or less tubular, somewhat swollen basally, and possesses spin- ules on the ventral surface. The distiphallus is connected to the sclerotized basiphallus by bph Fig. 2. Stevenia atramentaria (Meigen); aedeagus, lateral view. Abbreviations: aph = acrophallus, bph = basiphallus, d.pl = dorsal plate, dl.pr = dorsolateral processes, eph = epiphallus, spd.scl = spermduct sclerotization, v.pl = ventral plate. means of the dorsal plate, which divides distally into a pair of dorsolateral processes. The dorsal plate is extended ventrally on each side, forming two ventral plates. The acrophallus, carrying the phallotreme, is a simple, membraneous ex- tension of the distiphallus, probably encircling the three openings of the female spermathecal ducts during copulation. The aedeagus of many rhinophorids, e.g., Phyto spp. (figs. 15, 16), has not diverged markedly from this ancestral state, and the view is in agreement with that of Rikhter (1980), who mentions an epiphallus, basiphallus, a distiphal- lus immovably connected to the basiphallus, and “relatively” simple structure of distiphallic parts as the groundplan of the Tachinidae. Two features of the rhinophorid aedeagus de- serve mention. A possible autapomorphy for the Rhinophoridae is the well-developed ventral plates clearly set off from the dorsal plate and fused along the ventral margins, thus forming a sclerotized ring. Only the genus Paykullia pos- sesses unfused, but closely apposed, ventral plates, and this may be considered a reversal, as discussed below. It may seem somewhat odd to 20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 attach any importance to this character consid- ering the enormous variability of the distiphal- lus within the Tachinoidea, and certainly it is possible to enumerate several cases of non-rhi- nophorids (especially among the Calliphoridae) with fused ventral plates. However, most or all of these instances will be easily rejected as con- vergencies and I think the distinctive ventral plates will be of great value in the proper recog- nition of any rhinophorid. The other character to be mentioned is the sclerotization of the ventral part of the sperm- duct extending from the ventral plates to the phallotreme. In Phyto and Parazamimus this sclerotization is interrupted basally and does not reach the ventral plates (figs. 14—16). All other rhinophorid genera possess a sclerotiza- tion fused to the ventral plates and continuing to the phallotreme. The use of outgroup comparison for assessing the level at which this character is apomorphic is difficult to apply as the sister group of the Rhinophoridae is unknown. A similar sperm- duct sclerotization is of general occurrence in the Calliphoridae (the mesohypophallic sclero- tization of Salzer (1968)) but absent in most Sar- cophagidae and Tachinidae. If the interrupted spermduct sclerotization of Phyto is considered to be plesiomorphic within the Rhinophoridae then Phyto must be the sister group to all other genera. This hypothesis seems falsified by the several synapomorphies in the imaginal mor- phology of Phyto and Baniassa, and by the apo- morphic larval morphology of Phyto, which is also found in Paykullia and Melanophora. Probably the possession of a spermduct sclero- tization fused to the ventral plates is a ground- plan character in the rhinophoridae, and the spermduct sclerotization may be an important argument for a close affinity to the Calliphori- dae. To sum up, the characters which I regard as the most useful in the recognition of the family are the following: Larval characters: (a) cephalopharyngeal skeleton of first-stage larvae with toothed mandibles and elongated pharyngeal sclerite, (b) parasites of woodlice. Imaginal characters: (c) aedeagus with well-developed ventral plates that are fused (or closely apposed) along the ventral margins, (d) lower calypteres tongue-shaped, diverging from the scutellum, (e) metathoracic spiracle without a distinct o- perculum (except in Baniassa), (f) prosternum, proepisterna, greater ampullae, postalar walls, laterotergites, and suprasqua- mal ridges bare, (g) bend of vein M never greatly concave. It is important to note that the characters given not necessarily are rhinophorid autapo- morphies as some of them are found in other ta- chinoids as well. Character (f) is obviously ple- siomorphic within the Tachinoidea and is pro- vided to facilitate the exclusion of rhinophorid- like Calliphoridae. GENERA MISPLACED IN THE RHINOPHORIDAE The previous lack of an unambiguous defi- nition of the family has resulted in some moving about of a few genera. Crosskey (1977) in his review of the Rhinophoridae gives evidence for the exclusion of genera like Shannoniella Townsend (Tachinidae), Bezzimyia Townsend (Tachinidae), and Opsodexia Townsend (Calli- phoridae), all of which earlier have been consid- ered to belong to the Rhinophoridae (or to the Rhinophorinae as a subfamily of the Tachini- dae). This exclusion is accepted in the present paper and only the genera listed by Crosskey (1977), with the additions of Kugler (1978), will be treated in detail. Some of these clearly de- viate from the definition given above and ought to be excluded from the Rhinophoridae. Angioneura Brauer & Bergenstamm. Angioneura has long been treated as belong- ing to the Rhinophoridae, but North American authors, especially Downes (1955, 1965), have transferred it to the Calliphoridae, this view be- ing accepted by Wood (1979). Crosskey (1977) discusses this genus in the paragraph “included genera possibly not Rhinophoridae” but accepts its rhinophorid status. It is noteworthy that the genus Angioneura contains some species with enlarged lower calypteres, viz., A. obscura (Townsend), the only Nearctic species seen, and A. acerba (Meigen). The lower calypteres of the other species investigated, although distinctly diverging from the scutellum, are semicircular and not of the typical tongue-like shape charac- teristic of the Rhinophoridae. The larvae, still unknown from the first stage, seem to be parasites of snails rather than wood- lice. Two of the five Nearctic species of Angio- neura are recorded as having been bred from snails (Reinhard, 1929; Downes, 1965) and A. cyrtoneurina (Zetterstedt) from the Palaearc- Pave: Rhinophoridae 21 tic Region has been bred from the snail Succinea elegans Risso (Cepelak & Rozkosny, 1968). Bedding (1973) collected thousands of woodlice from about 50 localities in southern England in order to breed all native species of Rhinophori- dae. He did not, however, obtain any specimens of A. acerba or A. cyrtoneurina, the only Eng- lish representatives (Kloet & Hincks, 1976). On this evidence I find it highly unlikely that any species of Angioneura parasitizes woodlice. The presence of species with enlarged or semicircular lower calypteres, the life habit of the larvae as parasites in snails, and the ventral plates of the distiphallus which, although rather well-developed, are completely free of and widely removed from each other (fig. 3), clearly corroborate the exclusion of Angioneura from the Rhinophoridae, and I follow Downes (1965) in regarding Angioneura as a calliphorid. It is interesting that the exclusion of Angio- neura leaves the American continent without indigenous species of rhinophorids. Two spe- cies, however, have been established on this continent, both probably introduced from Eu- rope: Phyto discrepans, which occurs in south- ern Canada, and Melanophora roralis, which is recorded from southern Canada, the eastern United States, the West Indies (Jamaica, St. Thomas), and Brazil. Examined species: Angioneura acerba (Mei- gen, 1838), A. cyrtoneurina (Zetterstedt, 1859), A. fimbriata (Meigen, 1826), A. obscura (Townsend, 1919). Melanomya Rondani. This genus is apparently closely related to Angioneura, and Downes (1965) treats Angio- neura as a subgenus of Melanomya. No host re- cords are known for the single European spe- cies, Melanomya nana (Meigen), but as with Angioneura, the absence of any specimens of Melanomya nana in the material studied by Bedding (1973) reduces the probability of a woodlouse parasitizing habit. In addition, the ventral plates of the distiphallus are rather widely separated (fig. 4). The similarity to Angioneura will then indi- cate a position in the Calliphoridae. The metathoracic spiracle of M. nana differs from the typical, somewhat triangular, rhino- phorid type of spiracle (Crosskey, 1977: figs. 41—44) in being broad with a well-developed anterior fringe. This may provide further evi- dence for a calliphorid status as the majority of the Calliphoridae possess a rather large meta- thoracic spiracle, most often with a distinctly enlarged anterior lappet. Examined species: Melanomya nana (Mei- gen, 1826). Figs. 3—5. Aedeagus of Calliphoridae, lateral view: 3, Angioneura fimbriata (Meigen). 4, Melanomya nana (Meigen). 5, Morinia melanoptera (Fallén). A TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 Morinia Robineau-Desvoidy. This genus is accepted as belonging to the Rhinophoridae by Crosskey (1977) in spite of the presence of distinct hairs on the postalar walls, a character used by Crosskey to exclude rhinophorid-like Calliphoridae. Haired postalar walls occur in many Calliphoridae and in the subfamily Sarcophaginae of the Sarcophagidae (very seldom in subfamily Miltogramminae), but I have not found this trait in any tachinid or rhinophorid. The presence of haired postalar walls and the lack of well-developed ventral plates (fig. 5) make an inclusion under the Rhinophoridae somewhat improbable. Two other characters that may corroborate an exclusion are the well- developed metathoracic spiracular operculum (although an operculum is present in a single rhinophorid genus) and the presence of a weak- ly developed facial carina, these characters being most conspicious in the Japanese species M. ni- gerrima (Herting). A facial carina is not found in any rhinophorid but occurs freguently in the Calliphoridae and Tachinidae. On this sparse evidence I find a position in the Calliphoridae most corroborated. Examined species: Morinia melanoptera (Fallen, 1810), M. nigerrima (Herting, 1961). Termitoloemus Baranov. The only known species, T. marshalli Bara- nov, was originally described as belonging to the tribe Bengaliinae in the Calliphoridae. This was based on a similarity in life habits between Bengalia and Termitoloemus, predators of ants and termites, and similarities in the structure of the proboscis and palpi. Sabrosky & Crosskey (1970) transferred Termitoloemus to the Rhino- phoridae because of the possession of a simple metathoracic spiracle and tongue-like lower ca- lypteres. However, the lower calypteres of Ter- mitoloemus differ strikingly from all rhinopho- rids in having a distinct notch at the posterior base (fig. 6). The lappet of the metathoracic spi- racle is provided with stiff bristle-like hairs among the usual hairs. This condition is not found in the Rhinophoridae, but several groups of Calliphoridae possess stronger hairs on the anterior lappet. I have investigated the slide-mounted genita- lia of the male holotype of T. marshalli. The ae- deagus is highly apomorphic and very unlike that of any rhinophorid (or any other tachinoid) and its ventral plates are not fused (Baranov, 1936: fig. 1). This evidence, indeed, does not Fig. 6. Termitoloemus marshalli Baranov. Semidia- grammatical drawing of right lower calyptere of holo- type d. give much hint of the family affinity of Termito- loemus. The lower calypteres are not of the typ- ical tongue-like rhinophorid type but more sim- ‘ ilar to the plesiomorphic, enlarged type, and the metathoracic spiracle can be taken as evidence for either a calliphorid or a rhinophorid status. I do not find a rhinophorid assignment the most corroborative and I have chosen to consider Termitoloemus to belong to the Calliphoridae. Examined species: Termitoloemus marshalli Baranov, 1936. An inventory of the genera accepted as Rhi- nophoridae in the present paper is given in table 2. Note that Cirillia is treated as a junior syno- nym of Phyto. THE PHYLOGENY OF THE RHINOPHORID GENERA Very few attempts to create a suprageneric classification of the Rhinophoridae have been made, and these are often of little utility owing to the inclusion of several non-rhinophorid gen- era. Townsend (1935, 1938) divided his Mela- nophoridae (of which more than half of the gen- era were non-rhinophorids) into the five tribes Villeneuviellini, Melanophorini, Acampomin- thoini, Eggisopsini, and Moriniini. Séguy (1941), still with a rather broad (sub)family con- cept, arranged the few genera of which he had investigated the male genitalia into four groups based on perceived similarity. In the first group, Morinia (as Calobataemyia) is placed with Nyc- tia Panzer (Sarcophagidae), as Séguy apparently has dissected a specimen of Nyctia erroneously taken for a Morinia specimen (see his fig. 445, p. 343). Two other groups, both monogeneric, contain Stevenia and Melanomya (as Morinia), and the last group consists of Phyto, Rhinomo- Pare: Rhinophoridae 23 Table 2. Inventory of genera accepted as Rhinophoridae in the present paper. Following each generic name IS the number of species described at present (in brackets) and an indented list of species investigated in the present study. Acompomintho Villeneuve, 1927 (1 sp.) A. lobata Villeneuve, 1927 Azaisia Villeneuve, 1939 (2 spp.) A. obscura (Villeneuve, 1939) A. setitarsis Villeneuve, 1939 Baniassa Kugler, 1978 (2 spp.) B. fascipennis Kugler, 1978 B. paucipila Pape, 1985 Bequaertiana Curran, 1929 (2 spp.) B. argyriventris Curran, 1929 B. basilewskyi Peris, 1957 Callidesia Kugler, 1978 (1 sp.) C. pictipennis Kugler, 1978 Comoromyia Crosskey, 1977 (1 sp.) (C. griseithorax Crosskey, 1977; not seen) Macrotarsina Schiner, 1857 (1 sp.) M. longimana (Eggers, 1856) Melanomyoides Crosskey, 1977 (1 sp.) M. capensis (Zumpt, 1959) Melanophora Meigen, 1803 (2 spp.) Melanophora roralis (Linnaeus, 1758) Metoplisa Kugler, 1978 (1 sp.) M. carbonaria Kugler, 1978 Oplisa Rondani, 1862 (5 spp.) O. aterrima (Strobl, 1899) O. pollinosa Kugler, 1978 O. tergestina (Schiner, 1862) Parazamimus Verbeke, 1962 (1 sp.) P. congolensis Verbeke, 1962 Paykullia Robineau-Desvoidy, 1830 (8 spp.) P. brevicornis (Zetterstedt, 1844) rinia (as Metopisena), Angioneura, Rhinophora, and Melanophora. Herting (1961), in his revision of the Pal- aearctic species, divided the (sub)family into two tribes: Azaisiini (containing Azaisia and Acompomintho), with long antennae and elon- gate second aristal segment, and the clearly par- aphyletic Rhinophoriini, whithout these charac- ters. In the following is presented a phylogenetic analysis of the rhinophorid genera based on principles of phylogenetic systematics. Apo- morphies (numbers refer to the cladogram, fig. 30) are only given for genera with more than one species, as autapomorphies of single species (if present) are not necessary for cladogram construction. The species investigated are listed in table 2. As rhinophorids are sparse in museum collec- tions, most of the species were seen in only few (1—5) specimens. The monophyly of the Rhinophoridae, as de- P. kugleri (Herting, 1961) P. maculata (Fallen, 1820) Phyto Robineau-Desvoidy, 1830 (22 spp.) (Cirillia Rondani, 1856, syn. n.) P. angustifrons (Rondani, 1856) comb. n. P. cingulata (Zetterstedt, 1844) P. discrepans Pandellé, 1896 P. melanocephala (Meigen, 1824) P. pauciseta Herting, 1961 Queximyia Crosskey, 1977 (1 sp.) Q. flavipes Crosskey, 1977 Rhinomorinia Brauer & Bergenstamm, 1889 (12 spp.) R. capensis (Brauer & Bergenstamm, 1893) R. sarcophagina (Schiner, 1862) R. xanthocephala (Bezzi, 1908) Rhinophora Robineau-Desvoidy, 1830 (1 sp.) R. lepida (Meigen, 1824) Stevenia Robineau-Desvoidy, 1830 (18 spp.) S. angustifrons Villeneuve, 1913 S. atramentaria (Meigen, 1824) S. deceptoria (Loew, 1847) S. fernandezi Baez, 1978 S. hirtigena Herting, 1961 S. umbratica (Fallen, 1820) Tricogena Rondanı, 1856 (1 sp.) T. rubricosa (Meigen, 1824) Tromodesia Rondani, 1856 (2 spp.) T. angustifrons Kugler, 1978 Ventrops Crosskey, 1977 (> 1 sp.) V. milichioides Crosskey, 1977 V. spp. undescribed, Pape (in prep.) fined above, seems well corroborated by at least three synapomorphies: (1) cephalopharyngeal skeleton of first-stage larvae with toothed mandibles and an elon- gated pharyngeal sclerite, (2) parasites of woodlice, (3) distiphallus with well-developed ventral plates, which are fused along the ventral margins (secondarily free in Paykullia). Two monophyletic subgroups, the Phyto group and the Stevenia group, can be erected on larval morphology, as previously discussed. The monophyly of the Phyto group is corroborated by the apomorphy: (4) eighth abdominal segment of first-stage lar- vae with terminal lobes, a dorsal tongue, and paired ventral ridges. No shared apomorphic characters of the adult morphology have been found for the group, and as the first-stage larva is known for representa- tives of only three of the eight genera, the Phyto group is admittedly somewhat weakly founded. 24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 The first split in the Phyto group separates Pay- kullia + (Melanophora + Bequaertiana) from the remaining genera, this group possessing the synapomorphies: (5) female terminalia of the reduced non-teles- copic type, (6) wing cell r,,, long petiolate. Herting (1961) states that in the Palaearctic fauna only Paykullia and Melanophora possess shortened female terminalia (character 5), and Crosskey (1977), in his revision of the Afro- tropical fauna, notes that the female terminalia of the Afrotropical species apparently is of the normal telescopic type, although he did not dis- sect any specimen. Females of Bequaertiana are still unknown, but the assumed presence of non-telescopic ter- minalia seems well founded in the close affinity between Melanophora and Bequaertiana, as dis- cussed below. Character 6 is rather weak as the petiolate condition has arisen independently several times in the Rhinophoridae, and in Bequaertiana and Melanophora asetosa Kugler the bend of M is missing and an ancestral petiolate condition has to be assumed. Paykullia is a well-defined genus with the fol- lowing apomorphies: (7) distiphallus stout, possessing a strongly spinose pad on the ventral margin of each of the ventral plates and with the dorsal wall more or less prolonged (fig. 7), (8) male abdominal sternite 5 simple. As most male calyptrates possess a more or less excavated abdominal sternite 5, the simple, almost rectangular shape in Paykullia must be an apomorphic character. The monophyly of Bequaertiana + Melano- phora is corroborated by the synapomorphies: ( 9) parafrontalia with several (about 4—7) proclinate orbital setae, (10) male antennae with characteristic bottle- brush-like hairing (Crosskey, 1977: figs. 17 and 27). (11) hind coxae elongated. The hind coxae of Bequaertiana males (fe- males still unknown) are distinctly elongated; in both sexes of Melanophora roralis they are only slightly so. In addition, Bequaertiana and Mela- nophora possess very similar distiphalli (figs. 8, 9). The family affinities of Bequaertiana have been much discussed, Zumpt (1956) even sug- gesting an acalyptrate assignment. Crosskey (1977) doubts whether Bequaertiana is a rhino- phorid and although he notes the resemblance of the head to that of Melanophora roralis he is more inclined to accept a relation to Parazami- mus, another aberrant genus from the rainfo- rests of Zaire. The striking agreement in the apomorphic structure of the male antennae, the head, and the hind coxae of both Melanophora and Bequaertiana, however, leaves no doubt of their close affinity. Actually a case can be made for treating them as congeners. Melanophora asetosa, of which only the female is known, seems to be a typical Melanophora (as judged from the description in Kugler (1978)) except for the absence of the bent part of vein M, which is an apomorphic character of Bequaer- tiana! In the collection of the Zoological Mu- seum, University of Copenhagen, there is a sin- gle female Melanophora from Kenya, Naro Mo- ru, likewise with the bend of vein M missing. The terminalia appear to be of the short non- © telescopic type found in Melanophora and Pay- kullia (as seen in situ, the specimen is not dis- sected). On this evidence it seems most proba- ble (with a parsimonious concept) that the re- duced terminalia are a synapomorphy for the group Paykullia + (Melanophora + Bequaer- tana). The discovery of a female Bequaertiana and a male Melanophora asetosa may be most inter- esting, and if, as I think is most probable on the present evidence, the genus Melanophora is par- aphyletic with respect to Bequaertiana, it will be necessary either to fit Bequaertiana into the generic limits of Melanophora or to place M. a- setosa in the genus Bequaertiana. Melanophora asetosa and Bequaertiana share the apomorphy: (12) bent part of vein M absent (Kugler, 1978: fig. 15; Crosskey, 1977: fig. 28). eten wing venation occurs in Oplisa aterrima but is obviously a convergence. The genus Bequaertiana possesses some re- markable autapomorphies: (13) tibiae in males without clearly differ- entiated bristles, (14) male abdomen covered with thick silvery pollinosity, (15) wing vein R, strongly haired along its length. Melanophora (in the restricted sense with M. roralis as the only representative) is charac- terized by the distinctive white wing tips in fe- males. The sister group to Paykullia + (Melanopho- ra + Bequaertiana) is somewhat ill-defined and Pare: Rhinophoridae 25 10 Figs. 7—12. Aedeagus of Rhinophoridae, lateral view: 7, Paykullia maculata (Fallén). 8, Melanophora roralis (Linnaeus). 9, Bequaertiana argyriventris Curran. 10, Callidesia pictipennis Kugler. 11, Tromodesia angustifrons Kugler. 12, Baniassa fascipennis Kugler. may be polyphyletic. The possible monophyly of the group is corroborated by the single syna- pomorphy: (16) surstylar base extended medially (fig. 13). This may seem very conclusive, but several exceptions are found. The median extension is absent in Phyto pauciseta and both species of Baniassa, and indistinct in Phyto angustifrons. The first split in this group separates Tromode- sia + Callidesia from the remaining genera, their monophyly being corroborated by the synapomorphies: (17) clypeus distinctly bulging, (18) distiphallus of characteristic shape with the sclerotization of the spermduct bent dor- sally (figs. 10, 11). The two genera are depicted as sister groups on the cladogram (fig. 30), but they are very similar and could as well be treated as a single genus. I have not seen any specimen of Tromo- desia vibripennis Rondani, the type species of Tromodesia, and therefore I have not been able to. evaluate the monophyly of the genus, 1.e., to investigate whether 7. vibripennis is more 26 : TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 Fig. 13. Phyto melanocephala (Meigen). Cerci and surstyli, ventrolateral view, showing median exten- sions (m.ext) of surstyli. closely related to T. angustifrons than to any other species (or species group). The monophyly of the sister group of Tromo- desia + Callidesia seems well corroborated by the apomorphies: (19) lunula with setae, (20) notopleuron haired in addition to the usual two bristles, (21) katepimeron haired. All these traits occur sporadically in other rhinophorids, viz., many species of Paykullia and Rhinophora lepida possess some lunular se- tae; Tricogena, some Stevenia, and Rhinomori- nia sarcophagina may have a few additional no- topleural hairs; and some Rhinomorinia may have an occasional hair on the katepimeron (barette). However, the combination of these traits seems to have arisen only once in the Rhi- nophoridae. Baniassa is the possible sister group of Phyto and is well characterized by the apomorphies: (22) strongly holoptic eyes in males, (23) wings darkened apically, (24) wing cell r,,; petiolate, (25) metathoracic spiracle with operculum. As no other rhinophorids possess opercular metathoracic spiracles (character 25) this may be considered a reversal to the plesiomorphic condition. Baniassa paucipila Pape does not possess any of the synapomorphies 19—21 cor- roborating the monophyly of Baniassa + Phyto. However, the reduced hairing of Baniassa pau- cipila may be secondarily correlated with the yellow colouration of the thorax. Many yellow forms, e.g., the totally yellow species of Paraza- mimus and Bequaertiana from the rainforests of central Africa, possess a deviating, often re- Figs. 14—16. Aedeagus of Rhinophoridae, lateral view. 14, Parazamimus congolensis Verbeke; a = dorsal scle- rotization, dorsal view. 15, Phyto angustifrons (Rondani). 16, Phyto melanocephala (Meigen). Pare: Rhinophoridae 27 duced, hairing. This may be correlated with an association to a humid habitat. The aedeagus of B. fascipennis is shown in fig. 12. Phyto (including Cirillia) possesses the apo- morphies: (26) sclerotization of the spermduct interrupted (figs. 15, 16), (27) strong pre-alar bristle. Cirillia is characterized by the strongly devel- oped parafacial setae and a long-petiolate wing cell r,,;. These characters are likewise found in many species of Phyto, e.g., P. herting: Baez, and as Phyto does not possess any derived char- acters not shared with Cirıllia, a generic separa- tion between these seems unnatural in a phylo- genetic sense. Parazamimus is a strange monotypic genus from the tropical rainforests of Zaire. The single specimen known is in somewhat bad condition and the micropin, by which the head is mounted on the body, unfortunately penetrates the lunu- la, making it impossible to see whether setae are present. The structure of the distiphallus with the reduced sclerotization of the spermduct (fig. 14) is very reminiscent of Phyto, and Parazami- mus is tentatively placed as a sister group to Phyto although it does not possess any of the synapomorphies given for Baniassa + Phyto. Returning to the other group that could be erected on larval morphology, the Stevenia group, the possible monophyly is corroborated by the apomorphies: (28) setal bases of first-stage larvae produced into proleg-like structures, (29) acrophallus sclerotized and tripartite. Other genera like Parazamımus and Tromo- desia have the acrophallus partly sclerotized, but apparently developed independently and without the tripartition which is so characteris- tic of the Stevenia group. Typically the acro- phallus is divided into two lateral and one ven- tral sclerotization (the latter being the extension of the spermduct sclerotization), but often a dorsal acrophallic sclerite is more or less dis- tinct. In some genera this dorsal sclerite is sim- ple but in others it is provided with two lateral armlike processes. The three acrophallic scle- rites are more or less grooved and probably guide the sperm into the ducts of the female seminal receptacles; a functional analogue to the acrophallus of many Tachinidae and Sarcopha- gidae (for the latter see Lopes, 1966; Lopes & Kano, 1968). The first split in the Stevenia group separates Melanomyoides, Queximyia, Rhinomorinia, Rhinophora, and Ventrops from the remaining members of the group. All five genera have a general Rhinomorinia-like appearance and their monophyly is corroborated by the synapomor- phy: (30) dorsolateral processes of distiphallus fused into a single median sclerotization (fig. 22a). Queximyia is a monotypic genus from South Africa, easily recognized by the very long an- tennae and characteristic head profile (Cross- key, 1977: fig. 14). The possession of a strong pre-alar bristle suggests an affinity with Phyto, but a bare katepimeron, the lack of lunular se- tae, and the fusion of the dorsolateral processes of the distiphallus suggest this to be unlikely. The long antennae could be taken as evidence for a close affinity to either Azazsia or Acompo- mintho, but no other characters support this po- sition and the present assignment based on the aedeagal structure (fig. 18) seems the best cor- roborative. Ventrops is another well-defined Afrotropical genus, at present with only a single described species, but other species are known. The ae- deagus of V. milichioides is shown in fig. 17. Ventrops possesses the following apomorphies: (31) eyes greatly enlarged, occupying most of the side of the head and with a concave hind margin (Crosskey, 1977: fig. 13), (32) cerci very short and almost concealed be- tween the surstylar bases. The remaining three genera, Melanomyoides, Rhinomorinia, and Rhinophora seem to com- prise a monophyletic group corroborated by their apomorphic head structure: (33) epistome strongly warped (Crosskey, 1977: figs. 8—10, 12) Melanomyoides is a monotypic genus, its rep- resentative M. capensis being originally de- scribed as a species of Chaetostevenia Brauer (= Paykullia) by Zumpt (1959). Crosskey (1977) discusses the affinity of Melanomyoides to other (supposed) rhinophorid genera, and mentions an extreme superficial similarity to Melanomya and an even closer resemblance to Angioneura. These similarities, however, are founded in all three genera being composed of small, shining black flies with holoptic eyes in the male, characters which are not especially convincing; Crosskey concludes by stressing the resemblance in head profile and distiphallus between Melanomyoides and Rhinomorinia. Similarly, a case could be made for a sister forwards 28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 Figs. 17—22. Aedeagus of Rhinophoridae, lateral view: 17, Ventrops milichioides Crosskey. 18, Queximyia fla- vipes Crosskey. 19, Rhinophora lepida (Meigen). 20, Melanomyoides capensis (Zumpt). 21, Rhinomorinia xan- thocephala (Bezzi). 22, Rhinomorinia sarcophagina (Schiner); a = dorsal sclerotization, dorsal view. Pare: Rhinophoridae 29 group relation between Melanomyoides and Rhinophora, both having somewhat similar wings with a petiolate cell r,,;, but a very short petiole occurs in some Afrotropical Rhinomori- nia. Melanomyoides is easily distinguished by the almost leaflike surstyli, the holoptic male eyes, and the petiolate wing cell r,,.. The aedea- gus of M. capensis is shown in fig. 20. Rhinophora is likewise monotypic and is easi- ly separated from Melanomyoides by the di- choptic eyes in males and the presence of lunu- lar setae. The aedeagus is shown in fig. 19. The genus Rhinomorinia is difficult to char- acterize on external adult morphology and I have only found a single character which may establish the monophyly of the genus: (34) Distiphallus ventrally with a greatly en- larged spinous surface (figs. 21, 22). The long and slender cerci and surstyli (Crosskey, 1977: figs. 34, 35) may be another character, but a very similar condition is seen in Queximyia. The sister group to the four Rhinomorinia- like genera possesses the following apomor- phies: (35) acrophallus more complex, the sclerites being longer and more distinctly grooved, (36) dorsal wall of distiphallus extended. A dorsal extension is likewise found in Mela- nomyoides (fig. 20) but this is probably a con- vergence. Two other characters which may be synapo- morphies for this group are: (37) dorsal acrophallic sclerite well-developed, with two lateral arms, (38) hypandrium spoon-shaped. Character 37, however, is not found in Meto- plisa, most Oplisa and most Stevenia. Character 38 is especially distinct in Tricogena, Oplisa, Metoplisa, and Azaisia, and the flat hypandrium found in Stevenia must be secondarily derived. The first split in this group separates Acom- pomintho + (Azaisia + Macrotarsina) from the others. The monophyly of these three genera is corroborated by the following synapomorphy: (39) anal vein (A,) shortened. Acompomintho, the only genus endemic to the Oriental Region, is well defined by the long antennae with prolonged second aristal seg- ment, the well-developed parafacial setae (Lopes, 1938: pl. 1, fig. 2) and the long-petiolate wing cell r,,;. The aedeagus is shown in fig. 25. 0.2 mm i Figs. 23-25. Aedeagus of Rhinophoridae, lateral view: 23, Macrotarsina longimana (Eggers). 24, Azaisia obscura (Villeneuve). 25, Acompomintho lobata Villeneuve. Abbreviations: d.a.s = dorsal acrophallic sclerite; l.a.s = lateral acrophallic sclerite. 30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 Azaisia + Macrotarsina possess the synapo- morphies: (40) anterior katepimeral bristle much weaker than posterior one, (41) dorsal acrophallic sclerite well-developed, other acrophallic sclerites slender and situ- ated close together (figs. 23, 24), (42) gonopods (pregonites) thickened (only slightly in Azaisia). In Acompomintho (and all other rhinopho- rids) the two katepisternal (sternopleural) bris- tles (character 40) are subequal to equal in size. Macrotarsina is well characterized by the greatly prolonged male fore tarsi. Azaisia is more difficult to characterize; the most conspic- uous trait, which may be autapomorphic for Azaista, is: (43) antennae long, with prolonged second aris- tal segment. This character is likewise found in Acompo- mintho and may actually indicate a sister group relation between Acompomintho and Azaisia as accepted by Herting (1961), who established a separate tribe containing these two genera. However, I consider the sister group relation between Azaisia and Macrotarsina to be more corroborated by the present evidence. The monophyly of the sister group of Acom- pomintho + (Azaisia + Macrotarsina) is cor- roborated by the apomorphy: (44) ventral plates of aedeagus with a pair of processes, each supporting a spinous pad (figs. 26—29). The first split in this group separates Metopli- sa from Oplisa + (Stevenia + Tricogena). Kug- ler (1978), in his description of Metoplisa car- bonaria, mentioned the superficial similarity to Oplisa, but he erected the genus because the three humeral bristles of Metoplisa form an ob- tuse-angled triangle and not an almost right-an- gled triangle as in Oplisa. The latter configura- tion is used as a key character for the genus Oplisa by Herting (1961) and Kugler (1978), but both Stevenia (with S. hirtigena as an excep- tion) and Tricogena possess this character. As no other rhinophorids possess this arrangement of the humeral bristles, and as the arrangement in an obtuse-angled triangle is of widespread occurrence, the almost right-angled configura- tion is assumed to be a synapomorphy for Ste- venia, Tricogena, and Oplisa: (45) three humeral bristles forming an almost right-angled triangle. Oplisa was divided by Herting (1961) into the two subgenera Oplisa (as Hoplisa) sensu stricto, characterized by latero-reclinate ocellar bristles, and the monotypic Anoplisa with proclinate ocellar bristles. Kugler (1978) described two ad- ditional species of Oplisa, which both would fall into the subgenus Anoplisa, but as this is clearly a paraphyletic group (as defined by Herting) it is not accepted in the present paper. Oplisa is somewhat difficult to characterize by distinct autapomorphies. The enormously enlarged ejaculatory sclerite of O. tergestina, O. aterrima, and O. oldenbergi (Herting) (see Crosskey, 1977: fig. 40; Draber-Monko, 1978: fig. 18) is unique in the Rhinophoridae, but O. pollinosa possesses a normal-sized ejaculatory sclerite. The following apomorphies corroborate the monophyly of Oplisa: (46) distiphallus with the processes of the ven- tral plate, which support the spinous pads, situated on a stalked extension (fig. 27), (47) male cerci short and blunt, not separated apically, (48) surstyli broadened apically. It seems fairly corroborated that a sister group relation exists between Stevenia and Trı- cogena, which share the apomorphy: (49) parafacial plate with a row of strong setae. Both genera are very similar in external mor- phology and in the structure of the aedeagus (figs. 28, 29). Stevenia is a well-defined genus with the following apomorphies: (50) wing cell r,,; petiolate, (51) hypandrium flat, (52) mid femur in males with a posteroventral comb of short stout bristles apically. Some species do not, however, possess char- acter 52 (Herting, 1961), which may define an infrageneric subgroup. GENUS INCERTAE SEDIS Comoromyia Crosskey. Crosskey (1977) described the genus on a sin- gle female of C. griseithorax. I have not seen this specimen, which seems to be the only one known at present, and I have not been able to incorporate the genus into the cladogram on the basis of the description alone. Crosskey men- tions a possible relationship with Phyto, as Comoromyia possesses a strong pre-alar bristle, but the bare katepimeron weakens this argu- ment. I prefer to exclude Comoromyia from the cladogram (fig. 30) until more information is available, especially with regard to the structure of the aedeagus as this provides several of the set-defining characters of the present analysis. Pape: Rhinophoridae 31 ext ww ZO Sp.p Figs. 26—29. Aedeagus of Rhinophoridae, lateral view: 26, Metoplisa carbonaria Kugler. 27, Oplisa aterrima (Strobl). 28, Tricogena rubricosa (Meigen). 29, Stevenia atramentaria (Meigen). Abbreviations: ext = stalked extension of ventral plate; sp.p = spinous pad. ACKNOWLEDGEMENTS Centrale, Tervuren), R. W. Crosskey (British I am grateful to the following colleagues for Museum (Natural History), London), R. Dan- kindly providing me with a valuable material: ielsson (Zoologiska Institutionen, Lund), A. Drs. M. Baez (Universidad de la Laguna, Tene- Freidberg (Tel Aviv University, Tel Aviv), P. rife), E. de Coninck (Musée royal de l'Afrique Grootaert (Institut royal des Sciences Naturelle 32 de Belgique, Brussels), B. Herting (Staatliches Museum fiir Naturkunde, Stuttgart), H. J. Müller (Deutsches Entomologisches Institut, Eberswalde-Finow), K. A. Schmidt (American Museum of Natural History, New York), P. Tschorsnig (Staatliches Museum für Natur- kunde, Stuttgart), and N. E. Woodley (National Museum of Natural History, Washington, DE, I am indebted to Mr. R. W. Crosskey for his help during a visit at the Entomological Depart- ment of the British Museum (Natural History). Special thanks are extended to Dr. L. Lyneborg (Zoological Museum, Copenhagen) for valuable œ 9 = 5 ae a [= Ei 5 SN SRI “n E Sins DNS Oe iy oe COSE Se Ma EER Oo oe a © © >| D E ‚© N ~ e © zio een ra en ein. OVE wai at e x è * * 015 2, O14 924, 027 08 013 923, O22 O7 012 O18 O26 O17 O11 O21 010 920 09 O19 6 = 516 4 03 02 Fig. 30. Cladogram of the Rhinophoridae at the generic level. Numbers refer to apomorphies discussed in the TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 2, 1986 | help and advice during my study, and to Drs. V. | Michelsen and S. Andersen (Zoological Mu- | seum, Copenhagen) for many stimulating dis- cussions. Note added while this paper was already in press: The paper by H.-P. Tschorsnig, 1985, Die Struktur des mannlichen Postabdomens der Rhinophoridae (Diptera), Stuttg. beitr. Naturk. ser. A 375, pp. 1—18, appeared after submitting this paper for publication and a detailed dis- cussion of the hypotheses presented will appear in a future paper. 5 x que 5 2 8 È 2 = = eo 2 8 = 2 2 a £ 2 5 3 5 È beads > 5 ol ba Ene 5 3 Eo MME Nor So our = 2 Se NEO eo ES © Es si 3 0 ne e > NON AIS aeg ons S * è 032 052 048 034 031 043 051 550 647 I eee | 233 041 490 i 040 I ? 030 4 39 450 44 © Le 35 320 29 Lo 28 0 text. Genera recorded as woodlouse parasites are marked with an asterisk. | Pape: Rhinophoridae 33 REFERENCES Andersen, S., 1982. Revision of European species of Siphona Meigen (Diptera: Tachinidae). — Ent. scand. 13: 149—172. Andersen, S., 1983. 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Manual of myiology 2: 1—296, 9 pls. — Itaquaquecetuba, Sao Paulo. Townsend, C. H. T., 1938. Manual of myiology 4: 1—246. — Itaquaquecetuba, Sao Paulo. Verbeke, J., 1962. Contribution a l’etude des Tachini- dae africains (Diptera). — Explors hydro-biol. des lacs Kivu, Edouard et Albert 3(4): 81—185. Wood, D. M., 1979. Tachinidae and Rhinophoridae. Pp. 418—419 in Danks: Canada and its insect fau- na. — Mems ent. Soc. Can. 108. Zumpt, F., 1956. Calliphoridae (Diptera Cyclorrha- pha). Part I: Calliphorini and Chrysomyuni. — Explor. Parc natn. Albert. Miss. G. F. de Witte 87: 1—199. Zumpt, F., 1959. Diptera (Brachycera): Calliphoridae. — S. Afr. anim. life 6: 427—440. MEIN, RL: DEEL 129 AFLEVERING 3 1986 TS63 5 _ TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR Ù DE NEDERLANDSE ENTOMOLOGISCHE VEREBEGENG, INHOUD H. K. Prau. — Untersuchungen zur Konstruktion, Funktion und Evolution des Flugapparates der Libellen (Insecta, Odonata), pp. 35—123, figs. 1—30. ait voor Entomologie, deel 129, afl. 3 Gepubliceerd 10-X1-1986 LS 1e Je di 4 Lé à i î UNTERSUCHUNGEN ZUR KONSTRUKTION, FUNKTION UND EVOLUTION DES FLUGAPPARATES DER LIBELLEN (INSECTA, ODONATA) von HANS KLAUS PFAU à iy Institut fur Zoologie der Johannes Gutenberg-Universitat, 6500 Mainz, Deutschland (B.R.D.) Fa) ÄBSTRACT The sceletal morphology and mechanics of the flight apparatus of the Odonata are des- ===" | cribed. The different types of muscle systems are functionally interpreted, taking into ac- count various aspects of aerodynamics. — Electrophysiological experiments reveal a func- tional relationship between several mechanoreceptors located at the wing base (a chordono- tal organ and two rows of campaniform sensilla) and the pronation-supination movements of the wing. — A comparison of the flight apparatus of Odonata with those of Ephemero- ptera and Neoptera leads to the reconstruction of an ancient flight system in Pterygota. It may thus be concluded that flight ability has evolved only once, supporting the hypothesis of a monophyletic origin of the Pterygota. Within the Pterygota independent lineages have lead to highly autapomorphous characters in the flight apparatus of extant Odonata, Ephemeroptera and Neoptera. — The results of earlier research on functional morphology and evolution are discussed. INHALTSVERZEICHNIS Die Odonata (Libellen) sind stammesge- | schichtlich alte, palaeoptere Insekten, die relativ 35 Bu. unverändert bis ins Karbon zurückreichen Pee mule tun eg zer. Soe Re 35 (Hennig, 1969). Ihre stark autapomorphe | 1. Skelettmorphologie, Skelettmechanik und Pas macht eine systematische Einordnung M vie LA Ne 2 innerhalb der Pterygota schwer. Mehrere Mög- gene o SUR lichkeiten) wurden schon durchgespielt und Pkeloeie. ae men al 3 ; : Elugelantriebi( Eluemotorä)e a... Se 43 begründet (s. die Zusammenfassungen und kri- Drehbewegungen des Flügels um die Längs- tischen Kommentare von Hennig, 1969, Kris- ZIONS ors Wey es ea nee ea 46 tensen, 1975 und Matsuda, 1981). Dabei spielte Veränderung der Flügelschlagbahn — “Vor- die Beurteilung verschiedener Merkmale des und Zurückschwingen” gres Idales rear. 57 Flugapparates eine große Rolle. Matsuda (1981) 72. Elügelmechanorezeptoren................ 62 etwa postulierte eine polyphyletische Ent- Mechanische Beanspruchung der Rezepto- stehung der Pterygota (und unabhängige Ent- Bede IRE SISI do wicklung der Flugfahigkeit bei Odonaten einer- 3. Evolution der Flugapparate der Odonaten, SEE und Ephemeropteren A Neopteren ande- Ephemeropteren und Neopteren .......... 75 rerseits), wobei er ‚sich wesentlich auf 4. Diskussion und Ergänzungen ............. 91 Homologie- und Funktionsinterpretationen der Flügelmechanik, Muskelfunktionen und ae- grundlegenden Bearbeitung des Odonaten-Flug- rodynamischer Effekt... PEN 91 apparates von Tannert (1958) bezog. Er deutete Sensorische Kontrolle und Flugsteuerung ... 101 allerdings an, daß Schwierigkeiten existieren, Evolution ....... ATEN PE SPACE ER 104 die einem direkten Nachvollziehen (und Begrei- ni een im tabellarischen se fen) der Ergebnisse Tannert’s im Wege stehen: Danksagung. NEN 17 Wing articulation, especially that of Odonata, | Lennon rennes — Sun soe sade aun 117 is difficult to study. At one time, while working PDE ZU en MR RE MARE 121 on the Insect thorax (Matsuda, 1970), I gave up (Le Eee EE cio io an LS 122 studying wing articulation in Odonata after a LA EME A few hours attempt, and decided to rely comple- tely on the work by Tannert (1958), which must have been completed after years of study”. Oh- ne genaue Kenntnis und Bewertung der Struk- 36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 turen und Funktionen (die sich erst als Ergebnis komplizierter, wechselseitiger Erhellungen er- geben; vgl. Hennig, 1950, 1982) stehen stam- mesgeschichtliche Hypothesen aber auf einem schwachen Fundament. Das Beispiel der um- strittenen Evolution der Pterygota zeigt deut- lich, daß Ubereinstimmungen fast beliebig ent- weder als Konvergenzen, Symplesiomorphien oder Synapomorphien interpretiert und begrün- det werden können. Es ist aber für eine Ent- scheidung zwischen diesen Möglichkeiten not- wendig (wenn auch leider in vielen Fällen noch nicht möglich), Rekonstruktionen der Grund- pläne und der davon ausgehenden, natürlich auf jeder Stufe funktionsfähigen evolutiven Ab- wandlungen durchzuführen — unter Berück- sichtigung des Funktionszusammenhanges der Strukturen im ganzen “Apparat” und auch der ökologischen Wertigkeit des Apparates. Die Beurteilung der Merkmale des Odonatenflugap- parates darf schon deshalb keineswegs kom- mentarlos allein auf die Arbeit von Tannert gestützt werden, weil auch andere Autoren zu akzeptabel erscheinenden, im einzelnen aber ab- weichenden Ergebnissen kamen. Russenberger & Russenberger (1959/60) vermitteln z.B. in ih- rer (in der Literatur wenig beachteten) Arbeit einen umfassenden Einblick in die Konstruktion des Odonatenflugapparates; ihre Ergebnisse zur Mechanik des Skeletts wurden anschaulich do- kumentiert und mit Hilfe von Modellen unter- mauert. In der angelsächsischen Literatur sind vor allem die Arbeiten von Clark (1940), Nevil- le (1960) und Hatch (1966) zu nennen. Ver- gleicht man die Ergebnisse dieser Autoren untereinander, und mit den Befunden von Tan- nert, so findet man grundsätzliche Unterschiede (sowie auch Widersprüche und Unklarheiten in- nerhalb der einzelnen Arbeiten). Eine Entschei- dung zwischen den verschiedenen Hypothesen ist jedoch ohne eigene Anschauung und Bewer- tung nicht möglich. Diese Situation machte eine Neubearbeitung des Libellenflugapparates (der offensichtlich für die Klärung wesentlicher stammesgeschichtli- cher Fragen von Bedeutung ist) notwendig. Da- bei wurde erneut von einer Analyse der Thorax- und Flügelmechanik ausgegangen; v.a. dieser Weg bietet sich als der primäre Zugang zum Verständnis des Flugapparates an (Experimente an vor dem Windkanal fliegenden Libellen brin- gen dagegen erhebliche Schwierigkeiten mit sich — ıhre Interpretation wäre außerdem ohne die funktionsmorphologische Basis kaum möglich). Darauf aufbauend wurde der Versuch unter- nommen, die Funktion der Flugmuskeln (Kraft- wirkungen, Zeitpunkt der Kraftentwicklung, antagonistische Kraftbeziehungen) zu er- schließen, und — mit Hilfe (sicher vorläufiger) aerodynamischer Überlegungen — auch ihre Bedeutung für den Flug abzuschätzen. Die Analyse der Flügelmechanik stellte außerdem eine Voraussetzung für die Untersuchung me- chanorezeptiver, für die Flugsteuerung wesent- licher Sinnesorgane des Flügels dar. Die Ergeb- nisse zur mechanischen Beanspruchung und Funktion der Rezeptoren wurden elektrophy- siologisch und mit Hilfe von Funktionsmodel- len überprüft. Ein Hauptziel der Untersuchungen war die Erhellung der Evolution des Flugapparates der Odonaten — und auch der übrigen Pterygoten. Voraussetzung dafür ist eine vergleichende Analyse aller drei rezenten Teilgruppen der Pte- rygota, also der Odonaten, Ephemeropteren - und Neopteren. In der Literatur über die beiden letztgenannten Gruppen finden sich jedoch wie- derum v.a. morphologische Detailbeschreibun- gen; die Funktionsmorphologie der Flugappara- te wird (wie ım Falle der Odonata) keineswegs genügend berücksichtigt und nicht übereinstim- mend beurteilt. (Im Hinblick darauf unterliegt Matsuda (1981), der unsere Kenntnisse der außerordentlich komplizierten Flügelgelenkung der Ephemeroptera für ausreichend hält, einem Irrtum.) Auch für diese Gruppen mußte daher von eigenen Ergebnissen und Bewertungen aus- gegangen werden. 1. SKELETTMORPHOLOGIE, SKELETTMECHANIK UND MUSKELFUNKTIONEN MATERIAL UND METHODE Das Skelett- und Muskelsystem des Thorax wurde durch Sektion v.a. an großen Anisopte- ren (Aeshna cyanea Müll., Aeshna mixta Latr., Anax imperator Leach), zum Vergleich auch an verschiedenen Zygopteren und der Gattung Epiophlebia Calvert (als Vertreter der Anisozy- gopteren), untersucht. Parallel dazu wurden Mazerate herangezogen. Die Skelettmechanik kann nur an frischtoten Tieren analysiert werden (auch nach dem Ein- frieren in stark verdünntem Alkohol sind Libel- len noch lange fur Bewegungsuntersuchungen brauchbar). Zur Feststellung von Lage und Zu- sammenspiel der Gelenke wurden verschiedene Experimente durchgeführt: die Teile des Flügels wurden z.B. von außen bewegt; die Bewegun- Prau: Flugapparat der Libellen 37 gen wurden durch Zug an den Muskelsehnen (“von innen”) überprüft; einzelne Gelenke wur- den festgelegt oder durchtrennt; u.s.w. Zur Kontrolle wurden mechanische Modelle ange- fertigt (u.a. auch ein Gesamtmodell, das alle Be- wegungsmöglichkeiten enthält — Pfau, in Vorb.). Einige wesentliche Gelenk- oder Bie- gestellen der Kutikula (v.a. der Flügelbasis) lie- | gen so versteckt, daf eine direkte Beobachtung an intakten Strukturen unmôglich ist. In diesem Fall mußten die Teile freipräpariert, oder ganz aus dem Zusammenhang isoliert, untersucht werden (so mußte z.B. die Oberseite der Flügel- basis für eine gleichzeitige Sicht auf die mit wichtigen Gelenken versehene ventrale Kutiku- la gefenstert werden). Da jeder Eingriff auch zu Veränderungen der Mechanik führen kann, war es notwendig, an Einzelteilen gewonnene Er- | gebnisse jeweils an vollständigen Strukturen er- neut zu überprüfen. Die Flügelbasis erwies sich als kompliziert | (“verschachtelt”) gegliedert. Die Analyse der Mechanik wird z.T. dadurch erschwert, daß ei- ner Bewegung auch mehrere Drehachsen zu- grundeliegen können — daraus resultierende Verformungen von Teilen führen dann während des Bewegungsablaufs zur Veränderung der Achsen-Ausrichtung(en). Kaum eine Drehachse ist stabil. Übergeordnete Bewegungen (etwa der Flügelschlag) können die Lage von Achsen so verändern, daß andere (untergeordnete) Bewe- gungen in bestimmten Abschnitten des Schlags eingeschränkt oder vollständig gesperrt werden. Diese Komplikationen machen es notwendig, bei der Beschreibung und zeichnerischen Dar- stellung der Skelettmechanik starke Abstraktio- nen vorzunehmen. Die Muskeln des Flugapparates sind oft “zwi- schen” den verschiedenen Bewegungssystemen angeordnet, weisen also Hebelarme zu mehre- ren Drehachsen auf (“polyfunktionelle Mus- keln”). Aus der wechselseitigen Abhängigkeit der Systeme, oder aus ihrer mechanischen Be- grenzung, lassen sich in einzelnen Fällen Schlüs- se auf die zeitliche Einschaltung und Funktion eines Muskels ziehen. Da direkte Informationen über Kontraktionsstärke und -zeitpunkt fehlen (und die beim Flug wirkenden passıven Kräfte auch höchstens geschätzt werden können) mußte das Zusammenspiel der Kräfte in dieser Weise (hypothetisch) rekonstruiert werden. SKELETTMORPHOLOGIE In diesem Kapitel soll die Morphologie der thorakalen und pteralen Strukturen nur kurz um- rıssen werden — eine genauere Darstellung wei- terer Details wird in den folgenden Kapiteln vorgenommen. Bei der Beschreibung des Tergum wird hier zunächst v.a. das Mesotergum berücksichtigt. Während Vorder- und Hinterflügel (sowie Me- so- und Metapleurum) weitgehend gleich aufge- baut sind, unterscheiden sich Meso- und Meta- tergum stärker voneinander; darauf soll auf S. 61f. noch näher eingegangen werden. Der Auf- bau des Tergum wird nur für seine vordere bis mittlere Region dargestellt; kaudal-lateral ist das Tergum vielseitig biegbar (weicher) und “folgt” den verschiedenen Flügelbewegungen ohne wesentlichen mechanischen Einfluß (es ist dort also auch nicht hebelnd am Flügelschlag beteiligt). Das Sternum (höchstens als Ur- sprungsgebiet einiger hier behandelter Muskeln interessant) wird ganz weggelassen. (Detaillierte morphologische Beschreibungen finden sich in alteren Arbeiten; Zusammenfassung bei Matsu- da, 1970.) Bis auf wenige Ausnahmen werden nur sol- che Teile beschrieben und benannt, die als funk- tionelle Einheiten (= Bewegungseinheiten) zu erkennen sind. Da in der Beurteilung der Funk- tion (und der Abgrenzung von Funktionsein- heiten) wesentliche Unterschiede zu vorherge- henden Arbeiten bestehen, müssen zum Teil neue Bezeichnungen eingeführt werden. Einige ältere Homologievorstellungen konnten nicht bestätigt werden. Die fraglichen Strukturen (z.B. Pterale 1, Basalare) werden hier zunächst so neutral wie möglich benannt; auf ihre Ho- mologie wird v.a. im Kapitel 3 näher eingegan- gen. Tergum Auf der Höhe des Flügelvorderrandes befin- det sich ein stabiles vorderes Randelement des Tergum, die Tergalbrücke (Tb, Abb. 1—5). Sie verbindet die Costalplatten (CP, Abb. 1a und 3) des rechten und linken Flügels. Die Tergalbrük- ke steht auf beiden Seiten über ein Gelenk (t1) mit der Unterseite der dorsalen Wandung der vorderen Costalplatte (vCP, Abb. 3) ın Kon- takt. Von ihrer Mitte ragt ein unpaares Apodem ins Köperinnere, das Hebelapodem (HA, Abb. 1); es bildet Ansatz und Hebel (vgl. S. 58f.) der dorsalen Langsmuskeln (dlm, Abb. 1 und 2). An die Tergalbrücke schließt eine mittlere Region des Tergum (T, Abb. 1a und c) an. Diese ist etwa in der Segmentmitte auf beiden Seiten laterad zu je einem “Tergalzapfen” (TZ, Abb. la) ausgezogen, der zusammen mit einem da- 38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 acy Abb. 1. (a) Blick von dorsal-medial auf das Mesotergum und die Basis des rechten Vorderflügels einer Aeshni- de. Hinter der Tergalbrücke (Tb) ist das Segment durch einen Sagittalschnitt geöffnet, so daß Apodeme und Muskeln zu sehen sind. Membran fein punktiert — einige besonders verstärkte Membranstellen wurden hervor- gehoben: zwischen Tb und vTS (vgl. S. 60), zwischen CP und RS (vgl. S. 51) und zwischen TZ und RAP (bei t2). Dorsaler Ansatz des Chordotonalorgans (CH) schraffiert. TZ, G2 und t2 liegen in Wirklichkeit vertieft. (b) Schema des Vorschwingmechanismus des Vorderflügels (Ansicht wie in a). CP* und RAP zu soliden Platten vereinfacht, der Tergum-Mittenbereich (T in a) als 2-armiges Gestänge dargestellt. Bewegte Elemente schraf- fiert. (c) Tergale Mechanik — noch stärker vereinfacht: Das Hebelapodem (HA) wurde auf die Höhe des t1- Gelenks versetzt, wodurch der vom dlm genutzte Hebel zum Ausdruck kommt. Prau: Flugapparat der Libellen 39 Kort pas? a a," ae 9 A ip?) LE np fee gue? vc? Abb. 2. Flugmuskulatur. Blick von hinten-oben auf das mesothorakale Segment. Flügelbasis durchsichtig ge- dacht, Tergum weitgehend entfernt, Pleuralleiste und pleurale Gelenkköpfe dunkel hervorgehoben. Die Mus- keln wurden zur besseren Übersicht weit voneinander getrennt. Nur der dim ist paarig dargestellt. Linke Kör- perseite: Muskeln des Flugmotors (punktiert); Mitte und rechte Seite: Vor- Zurückschwingmuskeln (gekreuzt schraffiert) und pronatorisch-supinatorische Drehmuskeln (schraffiert). pa nur bei Zygopteren und Epiophlebia, dim im Metathorax der Anisoptera reduziert. Muskelfunktionen s. Tabelle 1, S. 112f. hinter liegenden Gelenksklerit (“2. Gelenkskle- rit” G2, Abb. 1a) eine komplizierte Gelenkstelle zwischen Tergum und Radioanalplatte (RAP, Abb. 1a) ausbildet. Zwischen Tergalzapfen und Tergalbrücke liegt, lateral von der Tergummittenregion, ein morphologisch komplexer Seitenbereich des Tergum. Hier ist (rechts und links) das Apodem des großen Dorsoventralmuskels (dvm1, Abb. la und 2) ins Körperinnere versenkt. (Aufer dem dvmi dient das Apodem auch den Muskeln dvm2 und tp sowie einem Tergocoxalmuskel als Ursprungs- bzw. Ansatzgebiet.) Die mediale Wand der doppelwandigen Einstülpung geht, von einer schmalen Membranzone (m) unter- brochen, in die Tergummittenregion (T) uber, die laterale Wand tritt (uber weitere Sklerite und Membranen) mit dem Fligel in Beziehung. Die Lateralwand wird hier als Tergalsklerit be- zeichnet. Dieser Sklerit ist bei Anisopteren im Metathorax einteilig, im Mesothorax dagegen durch ein Gelenk in zwei Teile gegliedert (vor- derer Tergalsklerit vTS und hinterer Tergalskle- rit RTS, Abb. 1a; vgl. auch S. 59f. — fur Zy- gopteren und Epiophlebia s. S. 109f.). Der vordere Tergalsklerit!) ist vorn gelenkig mit der Tergalbriicke verbunden (vgl. Abb. 1a und S. 60) und besitzt lateral ein in Tierlangs- richtung verlaufendes Gelenk zu einem hier als Randsklerit bezeichneten Seitenelement (RS, Abb. 1a und 3). Der untere Abschnitt des vor- deren Tergalsklerits bildet das Ursprungsgebiet des Tergopleuralmuskels (tp, Abb. 2 und 5). Der Randsklerit, der sich schmal zwischen dem vorderen Tergalsklerit und der Costalplatte in der Tierlangsrichtung erstreckt, ist vorn am Costalplatten-Tergalbrücken-Gelenk tl betei- ligt (Abb. 1a, 2 und 3); hinten ist er laterad zu einem in den Flügel hineinragenden Zipfel, dem Ansatz des vorderen Coxoalarmuskels (vca, Abb. 2 und 3), ausgezogen. Der hintere Tergalsklerit ist kaudal mit dem Tergalzapfen TZ verwachsen (Abb. 1a). Kurz davor vermittelt ein kleines Skelettelement, der 1) Das gut abgrenzbare Skelettelement wird in der Li- teratur oft mit dem Pterale 1 der neopteren Insek- ten homologisiert (vgl. Matsuda, 1970, l.c. S. 390). Dafür spricht jedoch weder seine Lage noch seine Muskulatur (s. auch S. 84 und 88). 40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 “1. Gelenksklerit” (G1, Abb. 1a), ebentalls zur Radioanalplatte. Direkt am lateralen Gelenk dieses Sklerits (dem Gelenk zur Radioanalplat- te) inseriert der hintere Coxoalarmuskel (hca, Abb. 2 und 3). Pleurum Die Pleuralleiste verzweigt sich am oberen Ende in die beiden Flügelgelenkköpfe, den vor- pan, pan, deren Gelenkkopf und das Fulcrum (vGK, F; Abb. 2 und 3). Diese Gelenkköpfe bilden, zu- sammen mit der Costal- bzw. Radioanalplatte, die pleuralen Hauptgelenke des Flügels (pl und p2; Abb. 7). Dicht bei der dorsalen Verzwei- gungsstelle der Pleuralleiste inseriert der Tergo- pleuralmuskel tp (Abb. 2 und 5), der weiter me- dial von der Seitenwand des dvm1-Apodems entspringt. Qin C Sc R- M Cr 3A x A, ( HZ] Ui ms 7 hca vca em eee een ee |; RAP fa sub2 sub3 Abb. 3. Schema der Basis des rechten Flügels einer Aeshnide (Blick von medial-dorsal). Die Membran zwischen den Skleriten wurde fast überall entfernt, die RAP dorsal aufpräpariert; dadurch werden Strukturen in der RAP (CH, fa) sowie die Flügelunterseite und das Pleurum (letztere eng schraffiert) sichtbar. Die Muskeln beider Drehbereiche des Flügels wurden eingezeichnet (bis auf den bei x, ansetzenden sub1, der v.a. als Senker zu be- trachten ist). x, kennzeichnet die Angriffsstelle des reinen direkten Senkers bas1. Die Ausrichtung des Flügel- Chordotonalorgans (CH) wurde (zur besseren Sicht auf das vordere Epifulcrum-Gelenk) geringfügig verändert (vgl. Abb. 1a). Prau: Flugapparat der Libellen 41 Der vordere Gelenkkopf ist eine Bildung, die nur den Odonaten zukommt (vgl. Kapitel 3). Die an dieser Stelle brückenartig verbundene Dorsal- und Ventralwandung der (auf dem vGK aufliegenden) Costalplatte (Abb. 7) ware damit als Analogie zur durchgangigen Sklerotisierung des Pterale 2 der Neopteren auf der Hohe des (hinteren!) Gelenkkopfes (Fulcrum) anzusehen. Die funktionelle Bedeutung der Stabilisierungen ist leicht einzusehen: in beiden Fallen entsteht durch Verschmelzung der Wandungen eine be- sonders verfestigte Auflagestelle des Flügels auf einem pleuralen Gelenkkopt. Die sog. Epipleurite, Basalare 1 und 2 und Subalare, die bei der Mehrzahl der Pterygoten im dorsalen Bereich des Pleurum liegen, sind bei Odonaten nicht ohne weiteres aufzufinden: Das Basalare 2 wird hier — in grundsätzlicher Ab- weichung zu bisherigen Auffassungen — als ein ursprünglicher, vorn in der Flügelbasis befindli- cher Teil des Flügels aufgefaßt; das Basalare 1 dagegen als eine pleurale Neubildung der Neo- pteren, die bei Odonaten (und Ephemeropte- ren) — noch nicht gelenkig abgesetzt — als fe- ster Bestandteil des Pleurum vorliegt (vgl. Kapi- tel 3). Der hinter dem Fulcrum liegende Epipleurit, das Subalare, ist bei Libellen als ein kleiner Sklerit im oberen Sehnenabschnitt des Muskels sub1 zu erkennen (es wurde hier nicht abgebildet); allerdings kann die Homologie ‚nicht gesichert werden (vgl. auch S. 42). Der Sklerit hat für die Mechanik des Flügels keine Bedeutung — er dient wohl v.a. der Verstär- kung der Sehne des großen Senkermuskels. Flügelbasis Die Fligelbasis ist aus zwei größeren Ele- menten, Costalplatte und Radioanalplatte!) (CP, RAP; Abb. 1a und 3), aufgebaut. Während eine der Radioanalplatte äußerlich ähnliche Sklerotisierung auch in der Flügelbasis der Ephemeropteren aufzufinden ist, bereitet die Homologisierung der Costalplatte (zumindest ihres proximalen Hauptteils) größere Schwie- rigkeiten. Verschiedene Autoren haben das Pro- blem dadurch gelöst, daß sie die Costalplatte insgesamt mit der Humeralplatte anderer Insek- ten homologisierten (s. z.B. Hennig, 1969, l.c. Abb. 25, oder Matsuda, 1970, 1979) und dem- zufolge annahmen, daß in der Flügelbasıs der Odonaten zwei Humeralplatten existieren. In der vorliegenden Arbeit wird nur die “distale 1) Radius-Analis-Platte bei Tannert (1958). Costalplatte” Tannert’s mit der Humeralplatte homologisiert (vgl. Kap. 3). Die abweichende bisherige Homologisierung der Costalplatte beruht wohl z.T. darauf, daß die vorderen Epipleurite, die Basalaria, bei Odonaten nicht (wie “üblich”) vorn unter der Flügelbasis anzutreffen sind. Einige Bearbeiter (z.B. Clark, 1940; Chao, 1953; Asahina, 1954) schlossen daraus, daß die große Kappensehne des Basalarmuskels als das ins Körperinnere versenkte Basalare zu betrachten ist. An die Kappensehne würde dann folgerichtig zum Flügel hin die Humeralplatte anschließen. Nach Matsuda (1981), der eine unabhängige Evolu- tion der Flugfähigkeit der Odonaten annimmt, fehlen die Epipleurite bei Libellen jedoch von vornherein (l.c. S. 391f.). In der Costalplatte (CP) können drei Teile unterschieden werden: vordere, mittlere und hintere Costalplatte (vCP, mCP und hCP; Abb. 3). Diese Bezeichnungen werden in Anlehnung an Tannert (1958) verwendet. Zur hinteren Costalplatte wird hier jedoch auch die “distale Costalplatte” Tannert’s gerechnet. Die hCP besteht damit aus der proximalen hinteren Co- stalplatte (phCP, Abb. 3; “regio posterior der proximalen Costalplatte” bei Tannert) und der distalen hinteren Costalplatte (dhCP, Abb. 3; “distale Costalplatte” bei Tannert; Humeral- platte bei anderen Autoren, vgl. Kap. 3). Diese Benennungsänderung soll die Darstellung er- leichtern und der engen funktionellen Be- ziehung von phCP und dhCP bei Pronations- Supinationsbewegungen (im Abschlagsdrehbe- reich, vgl. S. 47ff.) gerecht werden. Die Unterseite der mittleren Costalplatte bil- det, zusammen mit dem vorderen Gelenkkopf (vGK), das vordere pleurale Hauptgelenk pi des Flügels (Abb. 3 und 7). Wenig lateral davon inserieren der große, direkte 1. Basalarmuskel sowie der viel schwächere 2. Basalarmuskel (Abb. 2: bas1, bas2; Ansatzstelle des bas1 bei x, in Abb. 3). Die Unterseite der mittleren Costal- platte ist nach kaudal verlangert (dieser Ab- schnitt entspricht dem Basalare 2 der Neopte- ren, vgl. oben und Kap. 3) und tritt uber ein Gelenk mit der Radioanalplatte in Verbindung (c4, Abb. 3, 7, 9 und 10). Die übrigen, dorsalen Gelenke der Costalplatte werden weiter unten und auf den S. 48f., 58 beschrieben. Wahrend die Costalplatte sich distad nur in der Costa fortsetzt, stellt die Radioanalplatte (RAP) das Ausgangsgebiet der übrigen Flügel- langsadern dar. Sie ist (im Gegensatz zur CP) 42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 nicht scharf abzugrenzen, da ihre vorderen und hinteren Bereiche funktionell zum distalen Flügel (den “Flügelsektoren”, vgl. S. 43 und S. 46ff.) zu rechnen sind. Außerdem ist die RAP nicht (wie andere Autoren annahmen; vgl. Tabelle 1, S. 112f.) für sich gegenüber der CP beweglich. Der Begriff RAP umfaßt daher eigentlich keine Funktions- (= Bewegungs-) Einheit — er wurde dennoch hier zur Kenn- zeichnung eines morphologisch abgrenzbaren Flügelteils beibehalten. Wie bei Tannert (1958) wird auch die Sklerotisierung der Unterseite der Flügelbasis (kaudal von der ventralen mCP) zur RAP gerechnet. Die RAP stellt somit eigentlich keine Platte, sondern eine doppelwandige “Auf- treibung” der Flügelbasis dar. Durch sie kann man nach distal — wie bei einem Handschuh in die Finger — in die einzelnen Flügellängsadern gelangen (nach cranial ist der “Handschuh” zur CP hin offen, ebenso nach proximal-ventral zum Thorax-Lumen; Abb. 3). Die komplizierten Verformungen der RAP bei Drehbewegungen des Flügels um Längsach- sen (S. 46ff.) lassen innerhalb der RAP vier ver- schiedene Funktionsteile erkennen: 1. — In der Mitte der Unterseite der RAP liegt ein Sklerit, der vorn und hinten durch (gut “gängige”) Scharniergelenke (el, e2, Abb. 7) scharf begrenzt ist. Er wird hier als Epifulcrum (Ef, s. Abb. 3 und 7) bezeichnet. Verstar- kungsleisten der Innenwand des Epifulcrum (Abb. 10) sorgen dafür, daß der Sklerit nur schwer verformbar ist und so bei den Flügel- Verwindungsbewegungen (S. 46ff.) ein stabiles Widerlager bildet. Dieser wichtige (bisher an- scheinend noch nicht beschriebene und nur bei Odonaten abgrenzbare) Sklerit bildet proximal den Gelenkauflagepunkt des Flügels auf dem Fulcrum (p2). Wenig distal davon setzt der kräftige 1. Subalarmuskel (sub1, Abb. 2; An- satzstelle bei x, in Abb. 3) am Epifulcrum an. Er ist, wie der bas1, mit einer langen, hart skleroti- sierten Kappensehne versehen, die dorsal in eine membranöse Sehne übergeht. Im membranösen Teil der Sehne ist ein kleiner Sklerit, wahr- scheinlich der Rest des Subalare, zu erkennen (deutlich v.a. bei Zygopteren). Man könnte auch das Epifulcrum selbst mit dem Subalare homologisieren, das dann bei Odonaten in der Fligelbasis “inkorporiert” ware (vgl. auch die entsprechende Lage des hin- teren, zweiten Basalare im Flügel — s. S. 41 und Kap. 3). Da es jedoch nur schwer vorstell- bar ist, daß das Subalare die Stelle der ventralen Flügelbasis, die vorher dem Fulcrum auflag, verdrängt hat oder umgekehrt bei den übrigen Pterygoten sekundär aus dem Flügel ausgewan- dert ist (dann würde bei Odonaten ein nicht-ho- mologes Fulcrum/Flügelgelenk vorliegen), ist das Epifulcrum wohl eher als ein Teil der ursprünglichen ventralen, dem hinteren Gelenk- kopf aufliegenden Flügelbasis zu betrachten. 2. und 3. — An das Epifulcrum schließen vorn und hinten Seitenelemente der RAP an, die funktionell als Basisteile des distalen Flügels (der Flügelsektoren, in welche sie ohne gelenki- ge Abgrenzung übergehen; vgl. S. 43) aufzu- tassen sind. Sie werden als Costalsektor-Basis und Cubitalsektor-Basis bezeichnet (CoSB, CuSB; Abb. 7). Während die Unterseite dieser Teile (hinten bzw. vorn) mit dem Epifulcrum die schon beschriebenen Scharniergelenke el und e2 bildet, ist die Wand ihrer Oberseite in der RAP nicht deutlich abgegrenzt: die Costal- sektor-Basıs endet kaudal an einer (nur bei Be- wegung der Teile sichtbaren) Biegezone in der RAP-Oberseite, die dem vorderen Epifulcrum- Gelenk e1 der Flügelunterseite gegenüber liegt und ungefähr parallel zu ihm verläuft (vgl. S. 47ff.); die Cubitalsektor-Basis ist dorsal in komplizierter Weise über Biegestellen (die ebenfalls erst bei Bewegung erkennbar sind) “eingelenkt” (vgl. S. 53ff.). Nach vorn — zum Membranspalt zwischen RAP und CP (Abb. 1a) hin — ist die CoSB durch eine diagonal zur Costa ziehende, ader- ähnliche Verdickung (cr,, Abb. 3) begrenzt. Dort, wo diese auf die Costa trifft, besitzt die CoSB (bzw. die RAP) ein Gelenk zur (hinteren) Costalplatte (c3, Abb. 3). Die ventrale Wan- dung beider RAP-Seitenteile (CoSB und CuSB) ist auf der Innenseite mit Verstärkungsleisten versehen (Abb. 10), die — ähnlich wie die ent- sprechenden Bildungen des Epifulcrum — eine Verformung der Teile bei den Verwindungsbe- be (vel. S. 46ff.) verhindern. — Die verbleibende, dorsal zwischen er und CuSB liegende mittlere Zone der RAP wird hier — da sie nur schwer abzugren- zen ist (s. oben) — nicht gesondert bezeichnet. Außer den beiden schon erwähnten Basalar- muskeln (bas1 und bas2) steht noch ein weiterer Muskel mit der CP in einer Beziehung, der vor- dere Coxoalarmuskel vca. Er bewegt (indirekt, über den Randsklerit RS wırkend; vgl. S. 39 und 51) die hCP nach unten. Zur RAP ziehen (außer dem 1. Subalarmuskel) noch vier (bei Zy- gopteren und Epiophlebia fünf) weitere Mus- keln (Abb. 2 und 3). Auf diese soll auf S. 46ff. und 57ff. näher eingegangen werden. Prau: Flugapparat der Libellen 43 Fligel Distal von der Costal- und Radioanalplatte beginnt die eigentliche Flügelspreite. Sie besteht aus 2 Hauptteilen, die nach ihrer (proximalen) vorderen Randader benannt werden: Costalsek- tor und Cubitalsektor (CoS, CuS; Abb. 8). Der Costalsektor umfaßt die Fläche zwischen Costa und Radius (basal incl. Media) bis zum Nodus-Gelenk (n, Abb. 8). Diese Flache wird durch stabile Queradern zwischen Costa und Radius versteift (cr,, panl, pan2; Abb. 3). Pro- ximal-vorn tritt der CoS (über das Gelenk c3) mit der hinteren Costalplatte in Beziehung (vgl. S. 41f.), proximal-kaudal geht er (als CoSB) in die Radioanalplatte über. Der Cubitalsektor, der proximal den Cubitus (CuP: Cubitus posterior; s. Abb. 3 und 8) und die Analis (A) als Hauptadern enthält, ist über den Arculus (Arc), und andere Queradern bis hin zum Nodus, gelenkig mit dem Costalsektor verbunden. Seine Basis (CuSB) bildet, wie wei- ter oben beschrieben wurde, einen kaudalen Abschnitt der RAP. Innerhalb des Cubitalsek- tors kann kaudal noch ein (v.a. bei Anisopteren im Hinterflügel großer) Analsektor unterschie- den werden. Dieser Teil ist für sich anscheinend nicht aktiv beweglich. Er spielt wohl bei der Verwölbung des Flügels (bei pronatorischer Verwindung) eine Rolle (vgl. S. 49f. und S. 54). Die beiden Flügelsektoren sind in Wirklich- keit nicht scharf gegeneinander abgrenzbar. Sie gehen distal — da bei den Verwindungsbewe- gungen zur Flügelspitze hin mehr und mehr der ganze Flügel erfaßt wird — ineinander über. Je nachdem, ob der Flügel pronatorisch oder supi- natorisch verwunden wird, betrifft dies die mittleren und distalen Bereiche der Sektoren je- doch in unterschiedlicher, nicht symmetrischer Weise (vgl. S. 46ff.). FLÜGELANTRIEB (“FLUGMOTOR”) Der Flügel wird beim Auf- und Abschlag um die (durch die beiden pleuralen Hauptgelenke pl und p2 gebildete) Scharnierachse P1/P2 (Abb. 7) bewegt. Der dorsoventrale indirekte Hebermuskel dvm1 hebelt den Flügel (durch, Senkung des Tergum) nach oben, die direkten Senker basl und subl ziehen ihn nach unten (Abb. 2, 4 und 5). Libellen fliegen demnach mit einem “indi- rekt-direkten” Schlagmechanismus (vgl. z.B. Weber, 1933, Lc. S. 167; Pringle, 1957; Tannert, 1958). Neuerdings (vgl. z.B. Snodgrass, 1958; Nachtigall, 1968; Hennig, 1969, 1972; Hadorn & Wehner, 1974; Schneider & Günther, 1978) wird der Flügelschlagmechanismus der Odona- ten jedoch als Antagonismus direkter (!) Heber und direkter Senker beschrieben, und ausge- hend davon auch gefolgert, daß die Flügel der beiden Seiten unabhängig voneinander (also auch gegenläufig) geschlagen werden könnten. So dargestellt, und als Besonderheit der Libellen verallgemeinert, ıst dies falsch. Einerseits grei- fen die dvm1 nicht direkt am Flügel an; ande- rerseits sind rechter und linker Flügel durch die starre Tergalbrücke (Tb), an der die indirekten Hebermuskeln der beiden Seiten mittelbar an- greifen, gekoppelt (Abb. 1a, 2, 4 und 5). Eine gegenläufige Aktion der eng beieinanderliegen- den Hebermuskeln der beiden Körperseiten (in den Abb. wurden sıe aus Darstellungsgründen weiter auseinander gerückt) ist unwahrschein- lich und wurde bisher auch nicht nachgewiesen (vgl. dazu auch S. 45f. und Anm. 11, S. 115). (Entsprechendes betrifft natürlich auch die di- rekten Senker der beiden Seiten, welche die Ausgangsposition des Tergum, für einen erneu- ten Aufschlag, wiederherstellen.) Schließlich ist der Subalarmuskel, der, zurückgehend wohl auf Weber (1933), auch für Libellen als direkter He- ber verkannt wird (vgl. Schneider & Günther, 1978), mit Sicherheit ein Senker, so daß ein (uni- lateraler) Antagonismus direkter Basalar- und Subalarmuskeln ebenfalls nicht vorliegt. Die verbreitete Ansicht, daß Libellen einen “rein direkten” Flugmechanismus besitzen, geht z.T. anscheinend darauf zurück, daß medial der pleuralen Gelenkköpfe tatsächlich auch direkte Heber am Flügel angreifen; entsprechend ihrer geringeren Stärke können diese Muskeln jedoch nicht als Haupt-Antriebsmuskeln angesehen werden — es sind eindeutig Stellmuskeln mit akzessorischer Antriebsfunktion (vgl. S. 47ft. und S. 97f.). Die davon gut abgesetzten, weiter medial verlaufenden indirekten Heber (dvm1) sind viel machtiger und greifen deutlich am Ter- gum an; merkwürdigerweise werden sie oft nicht beachtet. Daß die Libellen wahrscheinlich phylogenetisch auf Vorfahren zurückgehen, die auch beim Abschlag einen indirekten Antriebs- muskel, den dorsalen Längsmuskel, einsetzten, wird auf S. 78ff. näher erläutert. Die Flügelschlagbahn verläuft — infolge der von hinten-unten nach vorn-oben geneigten Scharnierachse P1/P2 — nicht senkrecht, son- dern schräg, von hinten-oben nach vorn-unten. Der Flügelab- und -aufschlag ist also in Wirk- lichkeit ein Ab-Vor- und Auf-Zurückschlag. Die Anisopteren zeigen dabei gegenüber den 44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Tb t1 ae EN pi S S up li S E | dvm2 = bas1 Abb. 4. Schematische Darstellung der Funktion kleiner, tonischer Muskeln als zugfederartige “Drosselmus- keln” der Antriebs-Heber (linke Hälfte) und -Senker (rechte Hälfte). Querschnitt durch den Thorax auf der Höhe der Tergalbrücke. Zygopteren und Anisozygopteren (Epiophle- bia) eine steilere Ausrichtung dieser (festgeleg- ten) “Grund-Schlagbahnebene” (Abb. 28: großer Winkel À, in a) und b) gegenüber À in c); vgl. auch Tannert, 1958, lc. Abb. 37 und 38). Dementsprechend sind die Thoraxsegmente (und damit auch die Muskeln, Terga etc.) in den beiden Gruppen verschieden stark schrägge- stellt (in den Abbildungen wurde die schräge Ausrichtung der Segmente ın der Regel nicht berücksichtigt). Die Muskulatur des Flugapparates der Libel- len kann (relativ grob) in drei Kategorien unter- teilt werden: 1) Die mächtigen Auf- und Ab- schlagsmuskeln (die “Powermuskeln”) dvml, bas1 und sub1, die den Flügel als Ganzes bewe- gen; man kann sie als die wesentlichen An- triebsmuskeln eines übergeordneten “Flugmo- tors” ansehen. 2) Muskeln, die zwar auch den Flügelschlag und den Flügel als Ganzes betref- fen (weshalb sie ebenfalls dem Flugmotor zuzu- rechnen sind), aber — da sie erheblich schwä- cher als die Powermuskeln und wahrscheinlich tonisch aktıv sind — als “Einstellmuskeln des Flugmotors” betrachtet werden müssen (s. S. 45f.). 3) Muskeln, die in der Größe dazwischen liegen und die eigentlichen Stellmuskeln verkör- pern; sie verändern, mehr oder weniger unab- hängig vom Antriebssystem, durch Bewegung von untergeordneten Teilsystemen die Anstel- lung oder die Schlagbahn des Flügels (s. S. 46ff. Wel Se Shar) In vielen Fällen ist es allerdings noch fraglich, ob Muskeln tonisch oder phasisch aktiv sind (oder beide Kontraktionsmöglichkeiten besit- zen). Meine Versuche, dies durch elektrische Reizung festzustellen, führten nicht immer zu eindeutigen Ergebnissen. Einzelne Muskeln kontrahierten sich auf jeden Einzelreiz hin (dvm1, bas1, sub1, sub2, vca, hca, fa; auch beim dlm war dies in einem Experiment der Fall) — sie sind demnach als phasisch einzuschätzen. Andere Muskeln (bas2, sub3) sprachen auf Ein- zelreize nicht an und zeigten erst bei höherer Reizfrequenz eine Kontraktion; dies wurde als Anzeichen einer tonischen Aktivität angesehen. Prau: Flugapparat der Libellen 45 tp Tb ti BR / | | © A N f) Abb. 5. Schema des bistabilen Schlagmechanismus; Abschlag a > c, Aufschlag d — f. Querschnitte durch den Thorax auf der Höhe der Tergalbrücke. Die Pleuralleiste wird in der ersten Hälfte beider Schlagphasen (bis b bzw. e) seitwärts ausgelenkt; ihre Rückstellkraft ist durch den Tergopleuralmuskel (tp), der bis zur Schlagmitte gedehnt wird, einstellbar. Da eine gegenphasige Bewegung des rechten und linken Flügels unwahrscheinlich ist, muß der Flügel der an- deren Körperseite (mit gleicher Bewegungsrichtung) jeweils gedanklich ergänzt werden. Hier müssen weitere Untersuchungen, v.a. aber elektrophysiologische Ableit-Experimente an fliegenden Tieren, angeschlossen werden. Einstellmöglichkeiten des Flugmotors Zwei kleine Muskeln, der 2. Basalarmuskel und der 2. Dorsoventralmuskel (bas2, dvm2; Abb. 2) greifen mit sehr langen Sehnen (der eigentliche, die Muskelfasern enthaltende Teil ist also nur kurz) so am Flügel bzw. Tergum an, daß sie — wie die großen, dicht bei ihnen lie- genden Muskeln bast bzw. dvm 1 — als Ab- bzw. Aufschlagsmuskeln des Flugmotors be- trachtet werden könnten. Der Größenunter- schied zu den phasischen Antriebsmuskeln bas und dvm1 ist jedoch so gravierend, daß eine phasische Kontraktion (bas2 beim Abschlag, dvm2 beim Aufschlag) als relativ wirkungslos angesehen werden mufì (darauf wies bereits Hatch, 1966, hin). Nimmt man dagegen eine to- nische Kontraktion an!), so ergibt sich, daß der bas2 der Aufschlagswirkung des dvmi, der dvm2 dagegen der Abschlagswirkung des basl entgegenwirkt (Abb. 42). Damit könnten Flügelgeschwindigkeit und Amplitude beim Auf- und Abschlag getrennt eingestellt werden. Die Muskeln wären jetzt als Zugfedern (mit va- riabler Federkonstante!) jeweils in die Phase des antagonistischen Antriebsmuskels eingeschal- tet; sie wurden nur in dieser Phase “belastet”, in der anderen dagegen “entlastet”. (Dieser Ge- sichtspunkt wurde von Hatch, 1966 — und auch von Neville, 1960 — übersehen; s. dazu auch S. 92f. und S. 111ff. Zu dem funktionell ähnlichen Muskel sub3 vgl. S. 56.) Während die Muskeln bas2 und dvm2 jeweils nur in eine Schlagphase (drosselnd) eingreifen, kann ein weiterer, wohl ebenfalls tonischer Muskel, der Tergopleuralmuskel (tp, Abb. 2), den Flügelauf- und -abschlag (in symmetrischer Weise) beeinflussen. Der Muskel vermag die Pleuralleiste nach innen zu ziehen, was im Me- sothorax zu einer gegenseitigen Annäherung der Ränder eines dorsalen Membranspaltes, der sich zwischen den Episterna der beiden Körper- seiten befindet, führt. Drückt man bei einem frischtoten Tier die Pleuren nach medial, so schließt sich der Spalt ebenfalls, und die Flügel werden entweder auf- oder abgeschlagen — je nach ihrer Ausgangsstellung diesseits oder jen- seits der Schlagmitte. Da die starre Tb für einen konstanten Abstand zwischen den Gelenken tl der beiden Körperseiten sorgt, und der Abstand tl-p1 ebenfalls konstant ist, schwingen die Pleu- ren demnach beim naturlichen Auf- und Ab- schlag zunächst seitwärts (entgegengesetzt zur Kraftrichtung des tp: Offnen des Membran- spaltes) und dann, mit der tp-Kraft, nach innen zurück (Schließen des Spaltes). Das bedeutet, daß der tp (beim Flug tonisch kontrahiert) je- weils nur in der zweiten Schlagphasenhälfte als ') Am bas2 durchgeführte elektrische Reizversuche deuten darauf hin, daß zumindest dieser Muskel beim Flug tonisch kontrahiert ist (vgl. S. 44). 2) Ein weiterer dünner Muskel, der Tergocoxalmus- kel (hier nicht abgebildet; dvm5 bzw. tc in der Ta- belle 1, S. 112f.) könnte theoretisch eine ähnliche Funktion wie der dvm2 haben. Da er jedoch an die Coxa zieht, ist er wohl eher als Beinbeweger zu be- trachten. Im Gegensatz zum dvm2 ist die Sehne des dvm5 kurz, der eigentliche Muskelteil dagegen lang. 46 TIJDSCHRIET VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Synergist der Antriebsmuskeln der betreffenden Schlagphase wirksam ist; in der ersten Halfte beider Schlagphasen, in der die Pleuralwande nach lateral bewegt werden, ist er dagegen ihr Antagonist (vgl. Abb. 5). Der Tergopleuralmus- kel der Libellen ist damit als Einstellmuskel ei- nes bistabilen Mechanismus (“Klickmechanis- mus”) zu betrachten, ahnlich wie der Pleuro- sternalmuskel oder der Tergopleuralmuskel!) der Fliegen (vgl. Boettiger & Furshpan, 1952; Nachtigall & Wilson, 1967; Heide, 1971; Pfau, 1973; Pfau et al., 1977). Dieser Mechanismus bewirkt, daß die Antriebskrafte des Flugmotors beim Auf- und Abschlag unterschiedlich auf die beiden Schlagphasen-Hälften verteilt werden: sie werden anfangs (in der ersten Hälfte) teil- weise zur Überwindung der (durch den tp ver- änderlichen) “Pleuralfeder” eingesetzt, während ihnen in der zweiten Schlagphasen-Hälfte (jen- seits des instabilen Klickpunktes; Abb. 5b, e) die Federkraft des zurückschwingenden Pleu- rum wieder hinzugefügt wird (die zunächst “abgezweigten” Teilkräfte gehen dem Flü- gelschlag also nicht verloren). Eine Bedeutung des bistabilen Schlagmechanismus könnte daher darın liegen, daß die Antriebsmuskeln (die sich im Prinzip nur bis zum Klickpunkt zu kontra- hieren brauchen) im isometrischen Bereich ar- beiten können. Außerdem könnten Schlagfre- quenz und Thorax-Resonanzfrequenz aufeinan- der abgestimmt werden (entsprechend wie beim Pleurosternalmuskel der Fliegen, vgl. Nachtigall & Wilson, 1967). Der Flügelschlag kann also — abgesehen von Veränderungen der Kontraktionsstärke und Frequenz der Antriebsmuskeln — durch relativ schwache, tonische Neben- bzw. Verspannmus- keln beeinflußt werden. Die Wirkung der bei- den Muskeltypen (bas2, dvm 2 — tp) auf den Flügelschlag ist jedoch grundsätzlich verschie- den. Durch die Muskeln bas2 und dvm2 können wohl kleinere Schlagunterschiede zwischen rechtem und linkem Flügel bewirkt (oder aus- geglichen) werden. Bei symmetrischer Arbeit der Antriebsmuskeln sind damit Geschwindig- keits-, Phasen- und Amplitudenunterschiede möglich, die wahrscheinlich bei Steuerbewe- !) Dieser letztere Muskel, genauer der Pleuralleiste- Subtegula-Muskel (dessen medialer Ursprung bei verschiedenen Dipteren von der Subtegula zum Tergum-Seitenrand verlagert sein kann; vgl. Mickoleit, 1969, l.c. S. 160), ist wahrscheinlich — da der Sklerit vTS der Subtegula entspricht (vgl. S. 87f.) — dem tp der Odonaten homolog. gungen eine wesentliche Rolle spielen (vgl. auch S. 92f.). Die tp-Muskeln der beiden Körpersei- ten stellen dagegen die Stärke eines bistabilen Klickmechanismus ein. Dies setzt allerdings ei- ne weitgehend gleichartige (symmetrische) Schlagbewegung der Flügel beider Körperseiten voraus (Abb. 5), woraus geschlossen werden könnte, daß größere Rechts-Links-Unterschie- de der Flügelschwingung (bei Wendeaktionen) und ein stärkerer Einsatz des Klickmechanis- mus (beim schnellen Vorwärtsflug?) nicht zu vereinbaren sind. DREHBEWEGUNGEN DES FLUGELS UM DIE LANGSACHSE Die Bewegungsrichtungen des Flügels bei Drehung um die Langsachse sind durch die Be- griffe Pronation und Supination gekennzeich- net: Bei einer Pronation bewegt sich die Flügel- vorderkante nach unten und die Hinterkante | nach oben; dabei beschreibt die Hinterkante, infolge der (bei Insekten wohl in der Regel) weit vorn liegenden Drehachse, einen größeren Kreisbogen als die Vorderkante. Bei der Supina- tion wird die Vorderkante entsprechend nach oben, die Hinterkante nach unten bewegt. Pronation und Supination werden beim Li- bellenflügel durch zahlreiche Gelenke und ge- lenkahnliche Stellen in der Flügelbasis und im Flügel ermöglicht. Dabei kann der gesamte Drehspielraum in zwei Unterbereiche, “Ab- schlagsdrehbereich” und “Aufschlagsdrehbe- reich”, unterteilt werden (Abb. 6). Diese beiden Bereiche müssen getrennt betrachtet werden, da ihnen eine unterschiedliche Mechanik zugrunde liegt. Die Bezeichnungen für sie sind jedoch in- sofern nicht ganz treffend, als die Drehbereiche beim Flug den beiden Schlagphasen nicht genau “entsprechen” müssen. So kann der Flügel z.B. schon vor der oberen Schlagwende (also noch ın der Aufschlagsphase) in den Abschlagsdrehbe- reich hineinbewegt werden (vgl. S. 97f.). Die extremen Flügelanstellungen der beiden Berei- che (pa und So Abb. 6) werden jedoch si- cher nur innerhalb der jeweiligen Schlagphase erreicht. Die beiden Drehbereiche grenzen in einer mittleren Anstellung (0°, s. Abb. 6) aneinander. An dieser Stelle sind sie (von beiden Seiten her) durch Anschlage mechanisch getrennt. Wird der Flugel von der mittleren Anstellung aus pro- niert oder supiniert, so führen beide Bewegun- gen (also die Pronation im Abschlagsdrehbe- reich und die Supination im Aufschlagsdrehbe- reich) zu einer Verwindung des Flügels Prau: Flugapparat der Libellen 47 Abb. 6. Fligeldrehbereiche und -krafte (qualitatives Schema). Querschnitte durch den Flügel — das schwarze Dreieck kennzeichnet die Flügel-Vorder- kante und -Oberseite. Ab-Dr Abschlagsdrehbereich, Auf-Dr Aufschlagsdrehbereich (bei 0° durch An- schläge gegeneinander abgegrenzt); pp passiv-pronie- rende, ps passiv-supinierende Kräfte. (allerdings auf ganz verschiedene und nicht symmetrische Weise); dabei ist es gleichgültig, ob die Drehungen durch Muskelkräfte (aktıv, von proximal aus) oder durch Luftkräfte (pas- siv, von distal aus) bewirkt werden. Eine gleich- zeitige Aktivität von Muskeln, die den Flügel entgegengesetzt verwinden, kann als unwahr- scheinlich angesehen werden (jedenfalls inner- halb der Schlagphasen). Die den Flügel gleich- sinnig drehenden Muskeln der beiden Drehbe- reiche können andererseits aufgrund der mechanischen Trennung nur “hintereinanderge- schaltet” wirksam werden; z.B. kann der Pro- nator des Abschlagsdrehbereichs erst dann opti- mal “greifen”, wenn die supinatorische Verwin- dung im Aufschlagsdrehbereich vorher ganz rückgängig gemacht ist — der Einflußbereich des “rückdrehenden” Pronators im Aufschlags- drehbereich endet dabei beim 0°-Anschlag. So kann über die Grenze der Drehbereiche hinweg strenggenommen nicht von Synergisten oder Antagonisten gesprochen werden, selbst wenn der Flügel im Prinzip im gleichen oder entge- gengesetzten Sinn gedreht wird. Die verschiede- nen Muskeln sind demnach funktionell sowohl nach ihrem Drehbereich als auch nach ihrer Drehrichtung zu charakterisieren. Darüber hin- aus können anscheinend zwei Haupttypen von Drehmuskeln unterschieden werden: 1) “Wen- depunktsmuskeln”, die den Flügel am Auf/Ab- schlags- bzw. Ab/Aufschlagsumkehrpunkt dre- hen, und 2) “Einstellmuskeln” der Flügelanstel- lung, die die Flügelanstellung innerhalb der eigentlichen Ab- oder Aufschlagsphase beein- flussen. Die Drehbewegungen des Flügels um die Längsachse werden im folgenden “chronolo- gisch”, zunächst für den Flügel-Abschlag (Abschlagsdrehbereich) und dann für den Aufschlag (Aufschlagsdrehbereich), behandelt; dabei soll jeweils zuerst auf die zugrundeliegen- de Mechanik des Drehbereichs und dann auf die in ihm wirkenden Kräfte eingegangen werden. Abschlagsdrehbereich Mechanik. — Am Ende des Aufschlags geht die supinatorische Aufschlagsverwindung mit dem Ausschwingen des Flügels (bzw. seiner Verlangsamung durch die beginnende Kontrak- tion der Abschlagsmuskeln) zurück: passiver, v.a. durch elastische Rückstellkräfte bedingter Teil der Pronation der oberen Schlagwende ım Aufschlagsdrehbereich (vgl. S. 53ff). De Flügel vollzieht jetzt eine darüber hinausgehen- de Pronationsdrehung in den Abschlagsdrehbe- reich hinein, die mit großer Wahrscheinlichkeit aktiv, durch einen phasischen Muskel, verur- sacht wird (vgl. S. 50). Obwohl beide Vorgän- ge pronatorisch sind, müssen sie aufgrund der unterschiedlichen Mechanik der beiden Drehbe- reiche getrennt behandelt werden (s. oben). Im Abschlagsdrehbereich ist fast die gesamte Flugelbasis — die ganze Radioanalplatte (und mit ıhr die gesamte Flügelspreite) sowie ein Teil der Costalplatte — an der Drehbewegung betei- ligt (im Gegensatz zum Aufschlagsdrehbereich, in dem nur ein kaudaler Bereich der RAP be- wegt wird und die übrige Flügelbasis in Ruhe bleibt). Nur die mittlere und die vordere Co- stalplatte (mCP, vCP; Abb. 3) sind nicht (oder nur geringfügig, wie im Falle der vCP!)) betrof- fen; sie bilden damit — im Verein mit dem pleu- ralen, vorderen Gelenkkopf (vGK), auf dem sie aufliegen — das für die Bewegungen ım Ab- schlagsdrehbereich wesentliche Widerlager (dunkel hervorgehoben in Abb. 8, 9a, b und 14). Der bewegte Teil des Flügels soll hier, zur Ver- 1) Die vCP steht proximal mit der hCP in Verbin- dung und wird bei der Abbiegung der hCP (ss. 49) geringfügig verformt und um ti gedreht. Die- se Bewegung wurde hier — da kein Einfluß auf an- dere Bewegungssysteme ersichtlich ist — vernach- lässigt. 48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 N I c4 p2 NE I I LC CI P2/C4 Abb. 7. Schematische Darstellung einiger Teile der Flügelbasis sowie der Drehachsen des Abschlagsdrehbe- reichs. Blick von medial-dorsal auf die beiden Flügelbasisplatten, die auf der Hohe des Pleurum (und z.T. auch weiter distal) angeschnitten wurden; Membran weggelassen, Sklerite der Flügelunterseite schraffiert. Die phCP wurde ein-wandig (massiv) gezeichnet — sie stellt in Wirklichkeit eine doppelwandige Ausstülpung der Dorsal- seite dar, die kaudal in die Membran zwischen CP und RAP übergeht. Nur die Pronation wurde durch Bewe- gungspfeile (an den Achsen P2/C4, C1, E1 und am Gelenk c3) verdeutlicht; die Pfeile an der Achse P1/P2 (der durch die Gelenke pl und p2 gebildeten Scharnier-Schlagachse — pl stellt dabei für sich ein Scharniergelenk dar) symbolisieren den Flügelschlag. r pi einfachung der Beschreibung, als “Verstell- Drei Gelenke, die der Verstellflügel zu den flügel”!) bezeichnet werden. (nicht-mitbewegten) Widerlager-Teilen ausbil- det, bestimmen zwei Haupt-Drehachsen des !) Der Begriff wurde von Pfau & Honomichl (1979) are (ee. 7 une de DL auch für den bei Pronation bzw. Supination beweg- : ) a ur se Ci zieht durch vs ungefähr ten Teil des Lamellicornier-Flügels verwandt, soll In Flügellängsrichtung ausgerichtetes Biegege- aber, als reine Bezeichnung eines Funktionsteils, lenk zwischen der mCP und phCP (cl, Abb. 3 keine Homologievorstellung ausdriicken. Prau: Flugapparat der Libellen 49 und 7); diese Achse kann (in Annäherung; vgl. S. 50) als eine Scharnierachse aufgefaßt wer- den. 2) Die Hauptachse P2/C4 wird durch zwei Gelenkpunkte, die zusammen ein Scharnierge- lenk bilden, bestimmt: durch das pleurale Flù- gelgelenk p2 zwischen Fulcrum und RAP-Un- terseite (Abb. 7; das Gelenk ist gleichzeitig an der Bildung der Auf-Abschlags-Scharnierachse P1/P2 beteiligt — vgl. S. 43f.) und das weiter distal liegende (morphologisch komplizierte) Gelenk c4 zwischen dem ventralen Kaudalfort- satz der mittleren Costalplatte und der Unter- seite der RAP (vgl. S. 41; Abb. 3, 7 und 10; zu diesem Gelenk s. auch S. 57ff.). Die beiden Hauptdrehachsen kreuzen sich nach ihrem Austritt aus der Flügelbasis im Raum, ohne sich in einem Punkt zu schneiden. An der Bewegung des Verstellflügels ist, wie | schon dargestellt wurde, proximal nur der hin- tere Teil der Costalplatte (die hCP) beteiligt, die | Radioanalplatte dagegen als Ganzes. Von distal | betrachtet spaltet die Flügelfläche demnach bei | c3 (dem distalen Gelenk zwischen der CP und RAP bzw. CoSB; vgl. S. 42 und Abb. 3 und 7) nach vorn einen Basisteil (die hCP) ab und geht hinten als Ganzes in die RAP über. Beide T ile | bilden weiter proximal die oben beschriebenen Gelenke, die hCP das Gelenk c1 der Flügel- oberseite!), die RAP das Doppelgelenk p2/c4 der Unterseite. Wird der Verstellflügel um die Längsachse gedreht, so wird er zu gleicher Zeit in beiden basalen Gelenken bewegt; da beides Scharniergelenke mit unterschiedlicher Achsen- ausrichtung sind, wird er proximal unter Span- nung gesetzt und verformt (was weiter distal wiederum zu einer Verwindung der Flügelsprei- te fuhrt; s. unten). Die Ursache für die Verspannung der Flügel- basıs kann man sich leicht durch ein Experiment verdeutlichen: Trennt man die RAP von der hCP ab (durch einen Schnitt durch das Gelenk c3) und bewegt die hCP für sich (im Gelenk c1) — und anschließend die RAP für sich (im Ge- lenk p2/c4) — so beschreibt der Punkt c3 je- weils eine unterschiedliche Raumbahn; im einen Fall um die Achse C1, im anderen um P2/C4. Beim intakten Flügel existiert eine solche unab- hängige Beweglichkeit der Teile natürlich nicht und der Verstellflügel gerät bei einer Drehbe- wegung zwangsläufig unter Spannung. Die aus der Verspannung resultierende Ver- formung läuft in verschiedenen weiteren Gelen- ken und gelenkartigen Stellen ab. Abgesehen 1) Die phCP ist eine Vorwölbung der Flügeloberseite. vom tergalen Gelenk t2 (in dem sich die RAP gegenüber dem Tergum dreht) und weiter distal im Flügel liegenden Gelenken (s. unten), spielen v.a. zwei Gelenkstellen in der Flügelbasis eine wesentliche Rolle: 1) das schon erwähnte Ge- lenk c3, die Stelle, an der die Flügelfläche das vordere Basiselement hCP “abspaltet”; 2) das Gelenk e1 zwischen Epifulcrum und Costalsek- tor, ein vertieft liegendes und daher von außen schwer erkennbares Scharniergelenk der RAP- Ventralseite (vgl. S. 42). Die dem Gelenk el ungefähr gegenüberliegende Biegezone in der dorsalen Wandung der RAP (vgl. S. 42) wurde hier nicht gesondert benannt, da sie zusammen mit dem Gelenk e1 funktionell als ein Gelenk in der RAP aufgefaßt werden kann; Gelenk und Biegezone bilden gemeinsam das Basisgelenk des CoS, wobei das gut definierte Scharnierge- lenk ei für die Bewegungen des CoS maßgeb- lich ist. Eine pronatorische Drehbewegung des Flügels im Abschlagsdrehbereich (Abb. 7; Abb. 9a — b) kann jetzt genauer beschrieben wer- den — sie stellt eine gleichzeitige Bewegung der Flügelbasis in allen vier aufgeführten Gelenken dar: Die hCP wird um C1 nach dorsal gebogen (vgl. auch S. 50); die RAP bewegt sich zugleich um P2/C4, ihr Vorderrand geht nach unten; die RAP wird unter Spannung gesetzt, wobei ihr vorderer Teil, die CoSB, um die Achse E1 abge- bogen wird; hCP und RAP werden im Gelenk c3 gegeneinander verdrillt, was äußerlich daran zu erkennen ist, daß sich die vordere Costa- Kante vom Vorderrand der dhCP entfernt (Ver- größerung des durch Costa und dhCP gebilde- ten Winkels; vgl. Abb. 14 c — a). Der Ge- lenkpunkt c3 wird auf einer aus allen Bewe- gungskomponenten resultierenden Bahn bewegt. Für die Bewegung des Costalsektors ist, wie schon erwähnt wurde, die Ausrichtung der durch das Gelenk el festgelegten Costalsek- tor/Epifulerum-Scharnierachse E1 bestimmend. Diese Achse steht schräg zum Flügel und ver- läuft (bei horizontal gestelltem Flügel) von in- nen-vorn-unten nach auften-hinten-oben. Der Costalsektor (zusammen mit den am Nodus fest angekoppelten distalen Vorderrandadern; vgl. S. 54 und Abb. 8) wird daher bei einer Prona- tion nach ventral-kaudal, relativ zur in Ruhe bleibenden (“eingespannten”) Basis des Cus, bewegt. Diese Bewegung (die geringfügig ist und daher ganz proximal, bei el, kaum auffällt) führt in der basalen Flügelhälfte dazu, daß die Zwischenräume zwischen den vorderen Längs- 50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 adern zusammengedrückt und verschmälert werden (wobei Gelenkstellen zwischen den Quer- und Langsadern im Cubitalsektor und an der Grenze zum Costalsektor eine Rolle spie- len; vgl. Abb. 8) und bewirkt in diesem Bereich außerdem eine Verwölbung des Flügelprofils nach dorsal. Jenseits des Nodus wird dagegen mehr und mehr die mit dem CoS und den (dista- len) Flügelvorderrand-Adern verbundene ge- samte Spreite erfaßt und gegenüber dem basalen (stabileren, proximal eingespannten) Bereich des CuS verdreht (Verwindung der Flügelsprei- tel). Demnach findet bei einer Pronationsbewe- gung im Abschlagsdrehbereich eine pronatori- sche Drehung des ganzen Verstellflügels (um C1 + P2/C4) statt, der eine (ebenfalls pronato- rische!) Verwindungsbewegung überlagert ist — der Flügel wird dadurch distal stärker pro- niert als proximal. “Pronation als Ganzes” und “pronatorische Verwindung” gehen aufgrund des Zusammenspiels aller Gelenke zwangsläufig stets miteinander einher. (Zur aerodynamischen Bedeutung der Verwindung vgl. S. 96f.) Eine “eigentliche Drehachse” des Abschlagsdrehbe- reichs konnte daher in den Abb. 7 und 9 nicht eingezeichnet werden — sie entspricht weder C1 noch P2/C4, sondern verläuft (als “immate- rielle” Achse) zwischen ihnen im Raum. Da die hintere Costalplatte in Wirklichkeit über eine scharnierartige Biegezone (und kein echtes Scharniergelenk, wie hier zunächst vereinfa- chend beschrieben) mit der mittleren Costal- platte verbunden ist, ist die Achse C1 — und damit auch die im Raum verlaufende “eigentli- che Drehachse” des Flügels — darüber hinaus nicht ganz festgelegt (s. Abb. 9a, b). Alle be- schriebenen Achsen des Abschlagsdrehbereichs (C1, P2/C4, E1) verändern beim Flügelschlag (also bei der Bewegung des ganzen Flügels um die Schlagachse P1/P2; Abb. 7) ihre Ausrich- tung zum Körper. In Bezug auf den Flügel sınd sie jedoch entweder als stabil zu betrachten (P2/C4) oder machen eine dem Ablauf der Drehbewegung (nicht dem Flügelschlag- Ablauf!) fest zugeordnete Achsenverstellung durch (C1, E1; vgl. Abb. 9), so daß die Drehbe- wegungen also in einem vom Antriebssystem (“Motor”) mechanisch weitgehend unabhängi- gen System ablaufen (vgl. unten und S. 91ff.). Der Abschlagsdrehbereich ist relatıv klein; er ist zum pronatorischen Extrem hin (p,,,,, Abb. 6) wohl v.a. durch die (infolge der Verspannung und Verwolbung der Flügelfläche) zunehmend erschwerte Verwindung begrenzt. Dreht man den Flügel von dort (supinatorisch) zurück (Abb. 7, Pfeile jedoch in umgekehrter Richtung; Abb. 9b — a), so erreicht der Drehbereich bei 0° einen Anschlag, und zwar vor allem dadurch, daß die Costa bei c3 von unten auf die dhCP stößt (Abb. 14a — c). Kräfte. — Die Flügelfläche, die sich nach proximal in die hCP und RAP gewissermaßen “aufspaltet”, besitzt mehrere Hebel für prona- torische und supinatorische Muskeln (aktive Kräfte); die distale Flügelfläche selbst bildet den wesentlichen Hebel für die beim Flug angreifen- den, die Flügelanstellung von außen beeinflus- senden Luftkräfte (passive Kräfte). Nur ein Muskel, der hintere Coxoalarmuskel hca, greift an einem pronatorischen Hebel, dem nach vorn-proximal über die P2/C4-Achse vor- ragenden Randteil der RAP an (genauer Ansatz : s. S. 39f.; Abb. 2 und 3). Der Muskel wird daher hier als der wesentliche Pronationsmuskel der Auf-Abschlagswende angesehen (vgl. auch S. 97f.; Abb. 6 und 9a, b) — infolge seines Hebel- arms zur Schlagachse besitzt er außerdem eine Flügelheber-Funktion (vgl. S. 91f.). Der 1. Basalarmuskel bas1 kann dagegen (im Gegensatz zu bisherigen Auffassungen) als ein reiner Senker angesehen werden, da er an der mittleren Costalplatte (bei x,, Abb. 3 und 9a), also außerhalb der Verstellflügels (am Widerla- ger!) angreift. Die von einigen Autoren postu- lierte unabhängige Bewegungsmöglichkeit der beiden Flügelbasisplatten CP und RAP um die Schlagachse, die zur Erklärung einer Prona- tionsfunktion des basi herangezogen wurde (vgl. Tabelle 1, S. 112ff.), ist in Wirklichkeit nicht vorhanden: Da die CP an zwei Stellen (dorsal bei c3, ventral bei c4; vgl. auch S. 58) mit der RAP ın Kontakt steht, kann sie nıcht unabhängig von der RAP um P1/P2 bewegt und der Flügel damit auch nicht (durch eine Ab- schlagsbewegung der CP) proniert werden. Die CP nimmt die RAP bei einer Schlagbewegung stets einfach mit (wie auch umgekehrt die RAP die CP), ohne sie dabei um eine Längsachse zu kippen. Erst dadurch ist die im Gegensatz zu äl- teren Anschauungen vorliegende funktionelle Trennung der Flugmotor- und Flügelstell-Me- chanik gewährleistet (vgl. auch S. 92f. und S. 112ff.)! Der 2. Subalarmuskel sub2, der kaudal von P2/C4 an der RAP ansetzt, vermag den Flügel zu supinieren — im Unterschied zum sub3 (vgl. Prau: Flugapparat der Libellen 51 S. 56) jedoch höchstens bis zur 0°-Anstellung, der Grenze des Abschlagsdrehbereichs zum. Aufschlagsdrehbereich (Abb. 9b — a). Damit könnte der sub2 als ein Antagonist des hea an- gesehen werden. Wahrscheinlich steht der Mus- kel jedoch beim Abschlag (nach der hca-Kon- traktion) einer anderen Kraft antagonistisch ge- genüber, so daß seine Funktion nicht (zumindest nicht allein) in einer Einstellung der hca-Kraft zu sehen ist. Beim Abschlag kommen nämlich wesentliche passive Kräfte ins Spiel: In dieser Schlagphase wird die Flügelunterseite von der Luft angeströmt (Abb. 26a). Da die Flügeldrehachse des Abschlagsdrehbereichs vor der Flügelmitte (zwischen C1 und P2/C4; s. S. 50) liegt, und der größere Hebel sich demnach kaudal davon befindet, wird der Flügel beim Abschlag (nach der Pronation durch den hca) wie eine Windfahne passiv pronatorisch weiter gedreht. Der 2. Subalarmuskel kann nun die Flügelanstellung — gegen die pronierenden Luftkräfte — beim Abschlag so verändern, daß der aerodynamische Anstellwinkel vergrößert (der geometrische Anstellwinkel dagegen ver- kleinert) wird. Der Muskel verhindert damit, daß der Flügel im Extrem tangential und ohne Luftkrafterzeugung angeströmt wird; er be- wirkt, über die Vergrößerung des aerodynami- schen Anstellwinkels, eine Vergrößerung der erzeugten Luftkraft (vgl. Abb. 26a und S. 93ff.). Da der 2. Subalarmuskel lateral von der Schlag- achse an der RAP angreift, wird der Flügel gleichzeitig mit größerer Kraft abgeschlagen (zur Bedeutung dieser Doppelfunktion vgl. auch S. 92ff.). Auch der 1. Subalarmuskel sub1, ein Haupt- antriebsmuskel des Flügelschlags (Ansatz am Epifulcrum bei x,, Abb. 3 und 9a), besitzt einen supinatorischen Hebelarm zur Drehachse des Abschlagsdrehbereichs; er ist jedoch, verglichen mit dem Hebel des sub2, sehr klein. Da zu er- warten ist, daß bei einer stärkeren Kontraktion des sub1 mit der erhöhten Geschwindigkeit des Flügels auch die Geschwindigkeit der anströ- menden Luft vergrößert wird, und die passıve Pronation dadurch ebenfalls zunimmt, ist nicht sicher, daß die supinatorische Nebenfunktion des Muskels überhaupt äußerlich in Erschei- nung tritt (Abb. 26a). Neville (1960) sprach dem subl, nach Ausschaltexperimenten, eine Dreh- wirkung ab; er ging allerdings von einer ande- ren Flügelmechanik aus und übersah auch die durch die passive Pronation bedingte Täu- schungsmöglichkeit (vgl. Tabelle 1, S. 112f., und Anmerkung 15, S. 115). Auf die supinatorische Nebenfunktion des Muskels soll in der Diskus- sion (S. 92ff.) noch eingegangen wer- den. Bei Kontraktion des vorderen Coxoalarmus- kels vca wird der nach lateral in den Flügel ra- gende Fortsatz des Randsklerits (RS), und damit die über eine sehnenartige Zwischenmembran (s. Abb. 1a) mit ihm verbundene hintere Costal- platte, nach unten gezogen. Auch dieser Muskel vermag den Flügel demnach zu supinieren (Abb. 9b + a); da er proximal von der Schlag- achse P1/P2 ansetzt, wirkt er gleichzeitig Flü- gel-aufschlagend. Der Zeitpunkt der vca-Kon- traktion kann vorerst nur erschlossen werden: Beim Abschlag eingesetzt, würde der Muskel die Flügelgeschwindigkeit vermindern und den Flügel gleichzeitig supinieren. Diese Funktions- kombination erscheint für eine günstige Beein- flussung der Luftkrafterzeugung innerhalb der Abschlagsphase (im Gegensatz zur Abschlags- + Supinationswirkung der subi und sub2 — vgl. Diskussion S. 92ff.) nicht geeignet. Der Muskel vermag den Fligel andererseits nur im Abschlagsdrehbereich zu supinieren, nicht da- gegen im Aufschlagsdrehbereich. Das macht auch einen Einsatz während des Aufschlags un- wahrscheinlich. Da die supinatorische Verwin- dung des aufschlagenden Flügels (vgl. S. 53ff.) erst dann beginnen kann, wenn die pronatori- sche Verwindung des abschlagenden Flügels bis zum Anschlag rückgängig gemacht ist, bleibt als möglicher günstiger Wirkungsort das Ende des Abschlags bzw. der Beginn des Aufschlags (der untere Schlagumkehrpunkt)!). In diesem Fall bestünde der Vorteil, daf der vca — der (im Ge- gensatz zum sub2) infolge seines Aufschlag-He- belarms bis zum Abschlagsende gedehnt wird — im isometrischen Bereich arbeiten könnte; er wäre zu einer besonders effektiven (und wahr- scheinlich bis zum Anschlag des Drehbereichs bei 0° reichenden) Supination in der Lage. Je nach Kontraktions-Zeitpunkt und -Dauer wür- de der Flügel am Abschlagsende abgebremst und supiniert und/oder zu Beginn des Auf- schlags supiniert und beschleunigt (Einleitung bzw. Fortführung der Schlagumkehrbewe- gung). Auf eine mögliche zusätzliche Funktion des Muskels, die mit der hier postulierten 1) Auch Beobachtungen von Neville (1960), die allerdings anders interpretiert wurden (vgl. S. 99 f.), deuten auf eine phasische Kontraktion des Muskels am unteren Schlagumkehrpunkt hın. 52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 CoS phCP Abb. 8. Funktionelle Gliederung des Odonaten-Flügels (vgl. Abb. 9). Die CP (bis auf die dhCP schwarz) ist im hinteren Bereich (phCP) als “Spangengeriist” (durchsichtig) dargestellt; die RAP wurde dorsal großenteils ent- fernt, um die Teile der Flügelunterseite und das Pleurum zu zeigen. Die Wandungen der CoS- und CuS-Unter- seite (nur proximal sichtbar) wurden jeweils dichter punktiert als die der Oberseiten. IR;, Ry ,5, s. S. 110f. Prau: Flugapparat der Libellen 53 “Kontraktion zwischen Ab- und Aufschlags- phase” in Übereinstimmung steht, soll auf S. 60f. noch eingegangen werden. Aufschlagsdrehbereich Mechanik. — Am Ende des Abschlags wird der Flügel supiniert — einerseits passiv (Schlag- verlangsamung und dadurch verringerte Luft- anströmung), andererseits wahrscheinlich aktiv durch den vca (vgl. oben und S. 98). Der Muskel vca ware dabei sogar in zweierlei Hin- sicht ein Supinator: indirekt durch das Abbrem- sen des Abschlags (Verringerung der passiven Pronation) und direkt durch die Supinations- wirkung im Abschlagsdrehbereich. Der Flügel erreicht jetzt (ebenso wie am Ende des Auf- schlags) eine zwischen den Drehbereichen lie- gende Anstellung (0°, Abb. 6). Während des fol- genden Aufschlags kann er von hier aus nur im Aufschlagsdrehbereich, durch eine Bewegung des Cubitalsektors (CuS) relativ zum Costalsek- tor, supinatorisch weiter gedreht werden (der Abschlagsdrehbereich befindet sich — und bleibt — an seinem Anschlag). Auch diese Be- wegung, die aktiv oder passiv bewirkt werden kann (s. unten), führt zu einer Verwindung des Flügels. Im Gegensatz zur (pronatorischen) Verwindung des Abschlagsdrehbereichs, die mit einer Drehung des gesamten Verstellflügels ein- hergeht, wird der Flügel im Aufschlagsdrehbe- reich jedoch nur (supinatorisch) verwunden; proximal ist daran allein die Cubitalsektor-Basis (CuSB), der kaudale Teil der RAP, beteiligt. Die Cubitalsektor-Basis ist ventral mit dem kaudalen Rand des Epifulcrum über das Gelenk e2 (Abb. 7, 10 und 14) — ein Scharniergelenk, das für die Bewegungen im Aufschlagsdrehbe- reich als bestimmend angesehen werden kann — verbunden. Dorsal geht der Cubitalsektor über mehrere, komplizierte Biegestellen in die RAP über. Wie im Falle des Costalsektors (S. 42, 49) stehen sich also auch hier ein Scharnierge- lenk (in der ventralen Wandung der RAP) und eine, in diesem Fall komplexe, Biegezone (in der dorsalen Wand der RAP) gegenüber, die funk- tionell als ein einziges Basisgelenk des Flügel- sektors aufgefaßt werden können. Einzelheiten dazu sollen erst auf S. 54f., im Zusammenhang mit der Wirkungsweise des Muskels fa, be- schrieben werden. Die für die Cubitalsektor-Bewegung maß- gebliche Scharnierachse E2 (Abb. 9c, d und 15) ist schräg zum Flügel ausgerichtet — beim hori- zontal gestellten, in Schlagmitte befindlichen Flügel von innen-unten-hinten nach außen- oben-vorn. Der Cubitalsektor wird dement- sprechend (bei einer Supination) nach ventral- frontal-distal bewegt. Distal von einer aderfreien Zwischenzone zwischen den Längsadern R+M und CuP steht der Cubitalsektor vorn über die prägnante Arculus-Querader (Arc, Abb. 3 und 8) mit dem Costalsektor in gelenkiger Verbindung. Die an dieser Stelle vom Radius nach kaudal-distal (als Arculus) abbiegende Media-Ader kann nur in einem kurzen Gelenkgebiet (arc, Abb. 3) gegen- über dem Radius bewegt werden; die Media ist an dieser Stelle verschmälert, sie wird weiter proximal wieder dicker und verschmilzt dabei mit dem Radius zu einer Einheit. Distal vom Media-Radius-Gelenk arc tritt der Cubitalsek- tor uber mehrere Gelenke schwacherer Quer- adern und schließlich über das Nodus-Gelenk (n, Abb. 8) mit dem Costalsektor in Kontakt. Die CuS-Bewegung um E2 fuhrt — ebenso wie die Bewegung des CoS im Abschlagsdreh- bereich um El (s. S. 49f.) — zu einer Verfor- mung des Widerlager-bildenden anderen Sek- tors. Ausschlaggebend ist dabei, daf die Achse der ım Flügel vom Arculus bis zum Nodus an- einandergereihten Gelenke, die als ein einziges Scharniergelenk zwischen Cubitalsektor und Costalsektor aufgefaßt werden können, anders ausgerichtet ist als die proximale Drehachse E2 des Cubitalsektors. Eine Bewegung des Cubi- talsektors kann sich so nicht als eine einfache Schwenkbewegung gegenüber dem Costalsek- tor-Widerlager abspielen — sie führt zwangs- läufig auch zu einer Verformung des CoS: Der Costalsektor, der proximal als fest eingespannt betrachtet werden kann (seine supinatorische Bewegungsmöglichkeit um E1 ist mit Erreichen des c3-Gelenkanschlages erschöpft — vgl. S. 50), wird ım distalen Bereich nach dorsal ab- gebogen (die Aderstabilität nımmt nach distal ab). Außerdem wird auch der (gegenüber dem CoS weniger stabile) Cubitalsektor selbst ver- formt: in dem an den Arculus anschließenden (mittleren) Flügelabschnitt wird das Wellblech- profil zwischen den Längsadern verändert — mit fortschreitender supinatorischer Cubital- sektor-Bewegung werden die Winkel des Well- blechs spitzer, d.h. die Aderzwischenräume werden zusammengepreßt. Im Aufschlagsdrehbereich kommt es dem- nach bei einer Verwindung, wie im Abschlags- drehbereich, zu einer Verengung der auf den Arculus distal folgenden Aderzwischenräume, und auch — durch Verformung des Arculus selbst — zu einer Verkleinerung des Abstandes 54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 zwischen den proximal vom Arculus liegenden Adern R + M und CuP. Die Verwindungsvor- gange der beiden Drehbereiche sind jedoch in mehrerer Hinsicht nicht symmetrisch. Dem Abschlagsdrehbereich liegt nicht nur eine völlig unterschiedliche Mechanik zugrunde (vgl. S. 47tt.), die Verwindung ist in diesem Drehbe- reich außerdem viel geringfügiger als die (supi- natorische) Verwindung im Aufschlagsdrehbe- reich. Diese Asymmetrie wird durch die unter- schiedliche Stabilitat (und Form) der beiden Sektoren und durch die nicht-symmetrische Ausrichtung von E1 und E2 verstarkt. Weiter- hin spielen einzelne Gelenke entweder nur im einen oder nur im anderen Drehbereich eine Rolle: Arculus- und Nodus-Gelenk werden z.B. nur im Aufschlagsdrehbereich eingesetzt (s. unten); ein Gelenk der kaudalen Spitze des Flügeldreiecks (das bisher noch nicht erwähnt wurde) ermöglicht dagegen die nur im Ab- schlagsdrehbereich auftretende Wölbung des Flügelprofils (s. S. 50). Bei der Supination im Aufschlagsdrehbereich wird nach distal ein immer größerer Quer- schnitt des Flügels erfaßt und schließlich die gesamte Fläche bewegt, wobei v.a. der Gelenk- einschnitt des Nodus dem distalen Flügel- Hauptteil einen Spielraum gegenüber dem CoS gibt. Der geometrische Anstellwinkel (a, Abb. 26b) wird dadurch zur Flügelspitze hın allmählich größer (vgl. Abb. 27; zur aerodyna- mischen Bedeutung der Verwindung s. S. 96f.). Die Verwindbarkeit des Flügels im Auf- schlagsdrehbereich ist wohl v.a. durch die zu- nehmende proximale Verspannung der Spreite limitiert. Wird der Flügel vom supinatorischen Extrem (S,,,,) aus pronatorisch zurückgedreht, so endet der Aufschlagsdrehbereich (wie der Abschlagsdrehbereich) bei 0°; in diesem Fall v.a. deshalb, weil der Arculus (bei Planlage der beiden Sektoren) am Radius einen Anschlag er- reicht. Auch das Nodus-Gelenk läßt eine Cubi- talsektor-Bewegung über 0° hinaus nicht zu: der Spalt in der Vorderrandader schließt sich, wobei auch hier ein Anschlag gebildet ist, der ein erneutes Spalt-Offnen (bei der nun folgen- den pronatorischen Verwindung im Abschlags- drehbereich) verhindert. Kräfte. — Auch im Aufschlagsdrehbereich kann der geometrische Anstellwinkel des Flügels — in diesem Fall von der Cubitalsektor- Basıs (CuSB) aus — aktiv vergrößert oder ver- kleinert werden. Drei Muskeln setzen an der Cubitalsektor-Basis an (sub2, sub3, fa; Abb. 3), jedoch können nur zwei davon (sub3, fa) als zum Aufschlagsdrehbereich gehörige Stellmus- keln angesehen werden. Der Fulcroalarmuskel fa entspringt von einer kurzen Kappensehne, die an der Grenze zwi- schen Fulcrum und Epifulcrum beweglich auf- gehängt ist (Abb. 2, 3 und 8), und zieht, etwa in Körperlängsrichtung, zum kaudalen Ende der RAP. Dort greift der Muskel an der Kutikula der Oberseite der CuSB an. Da die Funktion des fa nur verständlich wird, wenn die skelett- mechanischen Verhältnisse parallel dazu genau beschrieben werden, soll das komplizierte dor- sale Gelenkgebiet der CuSB im Abschnitt “Kräfte” (und nicht im Abschnitt “Mechanik”) behandelt werden. Die Ansatzflache des fa stellt ein in die kau- dale RAP eingesenktes Gebiet dar, das nach ventral und lateral apodemartig etwas über den Rand der Vertiefung hinaus ins Körperinnere vorragt (Abb. 15). Nach kaudal-dorsal setzt es sich in einem Sklerit fort, der anfangs etwa senkrecht auf der Muskelansatzfläche steht, dann zur Postcubitus- und Analis-Basis hin um- biegt und in diese übergeht. Dieses geschwun- gene Skleritband, der Cubitalsektor-Hebel (CuSH, Abb. 1a, 3, 8 und 15), verbindet das Ansatzgebiet des Muskels außerdem mit der Unterseite der kaudalen RAP (die weiter vorn das Gelenk e2 zum Epifulcrum ausbildet); die Verbindungsstelle liegt da, wo die Dorsalseite des Flügels an der Basis der Analıs in die Ven- tralseite umschlägt. (Das Flügel-Ligament und die Membranula, die diese Stelle verdecken, wurden in den Abb. 3 und 8 weggelassen.) Der CuSH grenzt vorn an einen Membran- spalt (ms, Abb. 1a, 3 und 15), der proxımal von der Stelle ausgeht, wo der CuSH auf das An- satzgebiet des fa trifft. An dieser Stelle beginnt außerdem eine schmale Zone besonders harter (sich schwarz hervorhebender) Kutikula (z, Abb. 1a, 9c und 15), die sich nach frontal-dor- sal-distal erstreckt und an einem in Flügellängs- richtung verlaufenden Falz (f, Abb. 15) ge- lenkartig endet. Der Falz, der die RAP etwa auf der Höhe des Postcubitus durchzieht, kann als die vordere, dorsale Grenze der CuSB aufgefaßt werden: an dieser Linie “artikuliert” die CuSB (stark vereinfacht ausgedrückt) mit der RAP- Mittenregion, wobei sie v.a. durch die Sklerit- verstärkung z eine besondere Aufhängung er- hält. Die CuSB wird bei einer supinatorischen Verwindung des Flügels (vgl. S. 53) sowohl ventral (gegenüber dem Epifulcrum) als auch dorsal (gegenüber der vor dem Falz f liegenden Prau: Flugapparat der Libellen 55 RAP-Mittenregion) bewegt. Auf der Flugelun- terseite wird der Winkel zwischen den am Scharniergelenk e2 aneinandergrenzenden Tei- len verkleinert; auf der Oberseite wird das Ansatzgebiet des fa nach hinten(-oben) aus der RAP “herausbewegt” und der Muskel da- durch gedehnt (schematisch dargestellt in Abb. 9c > d, 14c + e und 15). Diese Bewegung des Muskelansatzgebietes verläuft sehr genau ın der Richtung der Muskelfasern des fa. Die Bie- gegelenke der dorsalen CuSB übersetzen dem- nach die Bewegung des Cubitalsektors um die Scharnierachse E2 in eine in der Muskelzugrich- tung verlaufende Bewegung des fa-Ansatzes. Beobachtet man den Fulcroalarmuskel (durch kleine Fenster in der Kutikula), so kann man bei supinatorischer Bewegung des Cubitalsektors deutlich die Dehnung des Muskels beobachten. Dabei wird der gesamte Muskel gedehnt (so- wohl die dorsalen wie auch die ventralen Faser- partien! Vgl. dagegen Tannert, 1958; Tabelle 1, S. 112f. und Anmerkung 6, S. 114). Aus dem Beschriebenen ergibt sich, daß der fa bei seiner Kontraktion der Supinationsbewe- gung des Cubitalsektors entgegenwirkt — er ist demnach ein Pronator des Aufschlagsdrehbe- reichs. Beim Aufschlag wird die Flügeloberseite an- geströmt. Da die maßgebliche Drehachse E2 der Cubitalsektorbewegung den Flügel so durchschneidet, daß der größte Teil seiner Fläche sich kaudal von E2 befindet, wird der Flügel beim Aufschlag passiv supinatorisch, zur Richtung der anströmenden Luft hin, gedreht. Die passiven Kräfte wirken also entsprechend wie beim Abschlag, in diesem Fall jedoch nicht pronierend, sondern supinierend (vgl. S. 51). Der aerodynamische Anstellwinkel kann dem- nach beim Aufschlag durch einen pronatori- schen Muskel vergrößert werden (vgl. Abb. 26b) — dafür kommt allein der fa, der einzige Pronator des Aufschlagsdrehbereichs, in Be- tracht (zur aerodynamischen Bedeutung des fa vel. S. 95f.). Der Cubitalsektor-Hebel CuSH ist als das wesentliche kraftübertragende Element zwi- schen dem Muskel fa und dem distalen Cubital- sektor zu betrachten. Er ist selbst biegbar und so funktionell weniger mit einem Hebel als mit einem flexiblen Kupplungsstück zu vergleichen. Je nach der Größe der Zugkräfte von distal (Luft) und proximal (fa) wird das geschwungene CuSH-Band verschieden stark abgeflacht (“in die Länge gezogen”), wobei der vor dem Skle- ritband liegende Membranspalt (ms) den für die verschiedenen Biegungszustände notwendigen Spielraum gibt. Für die Bewegung des CuSH sind aber noch weitere Strukturen ın der dorsa- len RAP, die hier nur kurz erwähnt werden können, wesentlich. So ist die RAP z.B. kaudal vom Gelenk t2 membranös eingeschnitten. Die- ser Spalt, der sich parallel zum medialen Rand der RAP erstreckt (Abb. 1a), spielt bei der Ver- formung der kaudalen RAP durch den fa eine wesentliche Rolle und hält sie außerdem an- scheinend von der Stelle t2 fern: Die Kraft des fa wirkt von kaudal auf den Membranspalt, wo- durch der RAP-Rand nach medial bewegt (ab- gespreizt) wird. Dadurch wird der Schub auf das tergale Flügelgelenk t2 und den Tergalzap- fen TZ verringert (die Bewegung des Randes nach medial wird ihrerseits in den Gelenken des Sklerits G2 “abgefangen”). Weitere strukturelle Besonderheiten der RAP sollen auf S. 62ff. be- schrieben werden. Das vordere Apodem des fa ist genau am pleuralen Hauptgelenk p2 des Flügels aufge- hängt (Abb. 3). Dadurch befindet sich der Mus- kelursprung, bezogen auf den Auf- und Ab- schlag des Flügels, in einer neutralen Lage, d.h., die Flügelschlagbewegung beeinflußt den Mus- kel an dieser Stelle nicht, der Muskel seinerseits nicht die Schlagbewegung. Anders verhält es sich mit der von der P1/P2-Achse weiter ent- fernt liegenden kaudalen Muskelansatzstelle; sie macht die Auf- und Abschlagsbewegungen mit. Da der Muskelursprung jedoch uber ein mem- branöses Sehnenstück am Fulcrum befestigt ist und so eine vielseitig bewegliche Aufhangung besitzt (die auch um die Achse P1/P2 drehbar ist — die Sehne wird dabei tordiert), fuhrt der Flugelschlag zu keiner Torsion des Muskels: der fa wird als Ganzes zusammen mit der RAP um P1/P2 bewegt und ist damit funktionell als flügelinterner Muskel zu betrachten. Er könnte sich theoretisch zu jedem Zeitpunkt des Flügelschlags kontrahieren. Beim Abschlag ist der Flügel jedoch proniert angestellt und wird außerdem so angeströmt, daß sich der Cubital- sektor auf jeden Fall an seinem pronatorischen Anschiag befindet (s. S. 51, 54). Der fa wird in dieser Phase nicht gedehnt (belastet) oder ge- staucht (entlastet), da er bei einer Veränderung der Flügelanstellung im Abschlagsdrehbereich zusammen mit der RAP bewegt wird, und der CuS dabei an seinem Anschlag bleibt. Nur die zur supinatorischen Verwindung des Flügels (und Abbiegung des CuS) führenden Kräfte be- wirken eine Dehnung des Muskels, dessen Funktion so auf den Aufschlagsdrehbereich ein- 56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 geengt werden kann. In diesem Bereich kann der fa den Cubitalsektor selbst bei maximaler Kontraktion (und fehlenden Gegenkraften) höchstens bis zu seinem pronatorischen An- schlag (0°-Anstellung, vgl. S. 46f., 54) bewegen — der Muskel ist wirkungslos, wenn der CuS durch andere Krafte an seinem Anschlag gehal- ten wird. Er könnte also auch tonisch sein (vgl. aber S. 44) und wäre dennoch nur beim Auf- schlag wirksam. Der 3. Subalarmuskel sub3 (Abb. 2 und 3) greift kaudal (wenig vor dem sub2) an der Un- terseite der Cubitalsektor-Basis an und zieht nach ventral-lateral-kaudal. Der Muskel ist rela- tiv schwach und kurz und enthält in seiner dor- salen Ansatzsehne ein langliches Stück Resilin. Da eine phasische Kontraktion des wenig lateral von der Schlagachse P1/P2 ansetzenden Mus- kels beim Abschlag nur von sehr geringer Wir- kung sein durfte, ist von vornherein anzuneh- men, daß der sub3 tonisch aktiv ist (vgl. z.B. Hatch, 1966; auch eigene elektrische Reizversu- ‘che — s. S. 44 — sprechen dafür). Seine Kraft- wirkung fällt dann aber (abgesehen von einer anfänglichen Federwirkung des Resilinstücks) nicht in die Abschlagsphase (wie Neville u.a. annimmt; vgl. Anmerkung 19, S. 116), sondern in die Aufschlagsphase: beim Abschlag wird der Muskel (wie eine Zugfeder) gestaucht und ent- lastet, beim Aufschlag dagegen belastet. Der sub3 würde demnach (ähnlich wie der bas2, vgl. S. 45) v.a. die Aufschlagsgeschwindigkeit des Flügels vermindern. Da er auch einen Hebelarm zur Cubitalsektor-Drehachse E2 besitzt!), supi- niert er den Flügel gleichzeitig (auf diese Dop- pelfunktion soll in der Diskussion noch einge- gangen werden — s. S. 95f; zur weiterhin möglichen Beteiligung des Muskels an der unte- ren Flügel-Wendepunktsdrehung s. S. 98). Möglicherweise ist die 2-tache Wirkung des Muskels z.B. bei sitzenden Libelluliden direkt zu beobachten: sie ziehen die Flügel, v.a. bei Annäherung eines Feindes, nach unten-vorn (manchmal mit wenigen, ruckartigen Bewegun- gen) und verwinden sie dabei stark supinato- risch. Die erste Schlagphase, ein Aufschlag, der die Libelle v.a. nach vorn treibt, ware damit “vorbereitet” (zur Vortriebswirkung des Aufschlags vgl. S. 96). Der 2. Subalarmuskel sub2 wurde schon auf S. 50f. besprochen und als der wesentliche Muskel zur Vergrößerung des aerodynamischen Anstellwinkels beim Abschlag interpretiert. Dieser Muskel setzt zwar wie der sub3 auch am Cubitalsektor an (Abb. 3), da seine Zugrichtung ~ jedoch praktisch mit der Ausrichtung des Cubi- talsektor-Epifulcrum-Gelenkes e2 zusammen- fallt, bewegt er die RAP supinatorisch als Gan- zes (was nur im Abschlagsdrehbereich möglich ist!). Er würde demnach — falls er beim Aufschlag kontrahiert würde — keine Supina- tionsbewegung des Cubitalsektors (relativ zum CoS) bewirken, sondern nur der Aufschlagsbe- wegung entgegenarbeiten. 1) Dieser Hebelarm ist im Metathorax der Anisopte- ren sehr klein — möglicherweise ein sekundärer Zustand, der im Zusammenhang mit der besonde- ren Spezialisierung des Segments in dieser Gruppe steht (vgl. S. 61f., 95 f., 101, 110). Abb. 9. Prinzipschemata der Flügelstellbewegungen. Die Teile wurden zu Platten (CP und RAP) und Ge- stangen (CoS, CuS) vereinfacht; die distalen Verbindungen der Flügelsektoren wurden weggelassen. Die RAP ist in den einzelnen Bildpaaren dorsal-proximal auf verschiedener Höhe angeschnitten; die CP zeigt im vorderen Bereich (vCP, mCP) einen Anschnitt auf der Höhe des Pleurum und wurde nur kaudal (phCP) in ganzer Aus- dehnung (nach proximal vorragend) gezeichnet. Widerlagerbildende (bei den Bewegungen ortsfeste) Teile dun- kel (Flügeloberseite) oder schwarz (Unterseite), ihre Anschnitte und Kanten weiß gekennzeichnet; bewegte Tei- le weiß (Flügeloberseite) oder schraffiert (Unterseite, Anschnitte und Kanten). Muskelbezeichnungen nur für kontrahierte Muskeln eingetragen. a — b Pronation im Abschlagsdrehbereich (b — a Supination). Die klei- nen Pfeile an der Achse C1 (die vereinfachend als Scharnierachse angesehen wird) deuten eine (geringe) Verstel- lung dieser Drehachse an (s. S. 50). Die “eigentliche” Drehachse liegt zwischen C1 und P2/C4 (s. S. 50). Der bei x, in (a) kaudal von der Achse P2/C4 (und lateral von der Schlagachse) angreifende subl wurde nicht einge- zeichnet (der bas1 greift bei x,, am Widerlager, an). Die dem vorderen Epifulcrum-Gelenk el (> Achse E1) gegenüberliegende Gelenkzone in der RAP-Dorsalseite (vgl. S. 42) wurde stark vereinfacht als Gelenkein- schnitt dargestellt. c > d Supination im Aufschlagsdrehbereich (d > c Pronation). Der sub3 wurde nicht eingezeichnet (er ist in d kontrahiert bzw. — bei passiver Supination — gestaucht). e > f Vorschwingen des Vorderflügels — der Flügel sei weitgehend abgeschlagen (f > e Zurückschwingen). Zur Wirkung des dim, dvm1 und pa s. S. 60f. (pa nur bei Zygopteren und Epiophlebia); zur Drehung der RAP im p2-Resilingelenk und Auslenkung von RAP+Fulcrum im Gelenk fu vgl. S. 58f. Prau: Flugapparat der Libellen 57 VERANDERUNG DER FLUGELSCHLAGBAHN — “VOR- UND ZURUCKSCHWINGEN” DES FLÜGELS Diese Bewegungsmöglichkeit des Flügels ist bei Anisopteren nur im Vorderflügel-Segment entwickelt, im Metathorax dagegen reduziert. Daß ein Vor-Zurückschwing-Mechanismus bei Libellen ursprünglich in beiden Segmenten vor- handen war (allerdings im Metathorax mit einer abweichenden tergalen Mechanik und einem “kontraren” Muskelantagonismus; s. S. 61f.), ergibt sich aus dem Vergleich mit Zygopteren und Anisozygopteren; auf diese Gruppen kann hier jedoch nur kurz eingegangen werden. 58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Mechanik. — Hier soll zunachst nur der Me- chanismus des Flügelvorschwingens beschrie- ben werden. Auf den Rückschwing-Vorgang (der sich, durch Umkehrung, daraus leicht er- gibt) wird im Zusammenhang mit den Kräften (s. weiter unten) eingegangen. Bei der Vorschwingbewegung bildet die Ter- galbrücke (Tb, Abb. 1a) zusammen mit der mittleren Tergalregion T und den beiden Flügeln (genauer den Flügeln ohne CP*1)) eine Bewegungskette. Sie beginnt bei der Tergalbrücke, die in ihren lateralen Ge- lenken ti gegenüber den CP* nach hinten geschwenkt wird (Abb. 1b, c; Bewegung um die quer zum Tier stehende Scharnierachse T1/T1). Dadurch wird die Tergalregion T nach kaudal verschoben, die beiden über die Gelenke t2 mit den Flügeln in Verbindung stehenden Tergal- zapfen (TZ) werden kaudad-dorsad gedrückt und bewegen gleichzeitig beide Flügel. Dabei spielt (jetzt für den einzelnen Flügel betrachtet) v.a. ein innerhalb des Flügels liegendes, aus zwei Gelenken zusammengesetztes Scharnier- gelenk c2/c4 (s. unten) eine wesentliche Rolle. Ein Flügel kann nämlich erst dann nach vorn schwingen, wenn die durch die beiden Gelenke c2 und c4 laufende Drehachse C2/C4 überhaupt “gebildet” ist und auch in einer bestimmten Ausrichtung zum Körper steht; da dies nur in einem kurzen, unteren Abschnitt des Flügel- schlags der Fall ist, ist der Mechanismus ın den übrigen Phasenabschnitten erschwert bis ge- sperrt. Die erste Voraussetzung ergibt sich aus der Lage und Struktur der beiden Gelenke c2 und c4: c2 liegt vorn in der Flügeloberseite zwischen der phCP und der dhCP (Abb. 1a und 3); c4 ist das schon bei der Besprechung des Abschlags- drehbereichs erwähnte Gelenk in der Unterseite des Flügels zwischen der CP (mCP) und der RUE (CAO Sa und 10) IDA €2 ombre Sica Gin Scharniergelenk darstellt, das bei Flugeldrehbe- wegungen im Abschlagsdrehbereich zusammen mit der hCP bewegt wird (wodurch sich die Ausrichtung der Achse C2 verändert), ist das zusammengesetzte Scharniergelenk c2/c4 (und damit die wesentliche Achse C2/C4 der Vorschwingbewegung) erst dann “funktions- fähig” wenn C2 auf c4 zielt. Dies ist der Fall, wenn sich der Flügel in einer bestimmten An- stellung, nämlich am supinatorischen Anschlag 1) Unter der CP* wird der proximale Teil der CP (vCP + mCP + phCP), also die CP ohne dhCP, verstanden (s. Abb. 1b). des Abschlagsdrehbereichs, befindet (dagegen steht die Achse C2 z.B. bei einem extrem pro- niert angestellten Flügel, bei dem die hCP stark nach dorsal gebogen ist, fast parallel zur Flügel- fläche, und die Vorschwingbewegung ist in die- sem Fall gesperrt; vgl. Abb. 9b, 14a und S. 47ff.)2). Das Gelenk c4, das sowohl im Ab- schlagsdrehbereich (bei der Bewegung des Flügels in den Gelenken p2/c4 und c1) als auch bei der Vorschwing-Bewegung (um C2/C4) ei- ne Rolle spielt, ist seiner Struktur nach beiden Funktionen angepafst (Abb. 10): die RAP greift (an der Stelle, wo das Epifulcrum-Gelenk el proximal beginnt) mit Fortsatzen so in die ven- trale mCP ein, daß Bewegungen um beide Drehachsen möglich sind; sie ist andererseits an dieser Stelle mit der CP so “verzahnt”, daf sich CP und RAP bei den Schlagbewegungen wech- selseitig “mitnehmen” (vgl. S. 50). Die zweite Voraussetzung folgt aus der sich im Verlauf des Flügelschlags relativ zum Körper verändernden Ausrichtung der Achse C2/C4. Diese steht-beim aufgeschlagenen Flügel etwa in, einer Sagittalebene (nahezu senkrecht, nach vorn geneigt), wird gegen Ende des Abschlags dagegen durch eine Horizontalebene bewegt. Während die Vorschwing-Bewegung beim auf- geschlagenen Flügel gesperrt ist — sie würde zu einer Bewegung des kaudalen RAP-Randes nach lateral führen, was jedoch infolge des seit- wärts nicht dehnbaren hinteren Tergalbereichs nicht möglich ist?) — wird sie im Verlauf des Abschlags, jenseits der Schlagmitte, allmählich “freigegeben”. Die Achse C2/C4 wird dabei an- scheinend mehr und mehr “entsprechend einer im Gelenk t2 bestehenden RAP-Bewe- gungsmöglichkeit relativ zum Tergum” ausge- richtet. Eine Vorschwingbewegung wäre jedoch dennoch nicht durchführbar, wenn nicht auch eine Bewegung gegenüber dem Pleurum statt- finden könnte; die Achse C2/C4 verläuft ja me- dial nicht durch das Gelenk p2 hindurch, son- dern wenig vor ihm vorbei (Abb. 9). Diese Re- lativbewegung der RAP wird wohl weitgehend 2) Somit kann eine Bewegung der dhCP im proxima- len Gelenk c2 erst dann stattfinden, wenn im dista- len Gelenk c3 der 0°-Anschlag erreicht ist (vgl. S. 50). Die nach Tannert (1958, lc. Abb. 29) be- stehende “unabhängige Wirkungsweise” der bei- den Gelenke c2 und c3 trifft also nicht zu. 3) Erst wenn man den kaudal von t2 befindlichen Rand der RAP vom Tergum vollständig abtrennt, ist die Bewegung um C2/C4 auch im oberen Schlagbereich möglich. Prau: Flugapparat der Libellen 59 \ AN C2/C4 \ P2/C4 Abb. 10. Das Gelenk c4 (Innenansicht — rechter Vorderflügel von Aeshna cyanea). Apodem des fa weggelassen. Punktiert: Membranverbindungen und Resilin (r); das Resilinpolster ist im Vorderflügel be- sonders groß und spielt dort auch beim Flügelvor-und -zurückschwingen eine Rolle (vgl. unten). Das Ge- lenk c4, das zwischen der ventralen mCP und der RAP liegt, gestattet Pronations-Supinationsbewe- gungen im Abschlagsdrehbereich (p, s; Achse P2/C4) und Vor-Zurückschwing-Bewegungen (v,z; Achse C2/C4); beim Flügelschlag sind RAP und (m)CP (durch die “Verzahnung” der Teile bei c4 — und durch ein weiteres Gelenk, c3) eng aneinander gekop- pelt. CH: ventraler Ansatzstift des Chordotonal- organs. durch das Resilinpolster des Gelenkes p2 selbst ermöglicht (Verlagerung und Drehung des Epi- fulcrum-Zapfens gegenüber dem Fulcrum), führt aber außerdem anscheinend — ın einem an der Gabelungsstelle der Pleuralleiste liegenden, ventralen Gelenk des Fulcrum (fu, Abb. 9e) — zu einer Seitwarts-Auslenkung des ganzen | Fulcrum. (Wahrscheinlich ist das Gelenk fu, das für eine federnde Auflage des Flügels sorgt, außerdem auch beim Flügelschlag von Bedeu- tung.) Die durch die Gelenke c2 und c4 verlaufende Drehachse C2/C4 steht schrag zum Flugel. Sie ist (beim horizontal gestellten Flügel) von un- ten-innen-hinten nach oben-außen-vorn ausge- richtet (Abb. 1b). Die relativ zur CP* stattfin- dende “Vorschwing”-Bewegung ist dement- sprechend kompliziert: die Flügelspitze bewegt sich zunächst v.a. nach vorn-unten und dann — da die flügelinterne Achse C2/C4 zusammen mit dem gleichzeitig weiter abschlagenden Flügel weiterbewegt wird und so ihre Ausrich- tung verändert — nach medial-dorsal (Abb. 28, 29), wobei gleichzeitig auch eine Art “Supina- tion” stattfindet. Diese letztgenannten Bewe- gungen sind jedoch als Komponenten der Vorschwing-Bewegung anzusehen und dürfen nicht mit den auf S. 46ff. und 43 ff. beschriebenen Drehbewegungen um die Längsachse oder mit der Schlagbewegung des Flügels (beide gesche- hen ja um andere, eigene Achsen) verwechselt werden. Man könnte die Vorschwing-Bewe- gung aufgrund ihrer supinatorischen Kompo- nente (die am kaudalen Senken der RAP in der Abb. 1b und 9f zu erkennen ist) auch als eine Art “Fortsetzung” der Supination des Ab- schlagsdrehbereichs mit anderer Mechanik (und Muskulatur) bezeichnen: sie beginnt erst dann, wenn sich der Abschlagsdrehbereich an seinem Anschlag befindet (s.S. 50 und weiter oben). In der Bewegungskette Tb-T-RAP wird eine Schwenkbewegung der Tergalbrücke (um die Querachse T1/T1) und Schubbewegung des Tergum in eine Flugelbewegung (um die schrag zu T1/T1 stehende Achse C2/C4) “umgesetzt”. Dies ist nur möglich, wenn noch weitere Gelen- ke und Biegestellen mitwirken. So spielt z.B. ein Gelenk zwischen der Tergalbrücke und der mittleren Tergalregion T (g, Abb. 1b, c) eine wichtige Rolle. Außerdem ist der Tergalzapfen TZ gegenüber T biegbar — er wird in einem Gelenkeinschnitt seiner Basıs tordiert (der Ge- lenkeinschnitt wird während des Schubvorgan- ges verengt). Schließlich wird eine Bewegung des Tergum nach kaudal, relativ zur CP*, v.a. dadurch ermöglicht, daf die Tergum-Mittenre- gion T gegenüber den tergalen Seitenbereichen in der Tierlängsrichtung verschiebbar ist (s. auch weiter unten): die Bewegung läuft proxi- mal an der CP* entlang, “umgeht” sie gewisser- maßen. Da das Gelenk pl der CP* ein in der Tierlangsrichtung verlaufendes Scharniergelenk darstellt (vgl. z.B. Abb. 7), ist andererseits distal dafür gesorgt, daß die CP* das für eine effektive Vorschwingbewegung notwendige stabile Wi- derlager bildet; die Bewegung wird dadurch von dem die Costalplatte betreffenden Teil des Flügelantriebs unabhängig. Wie auf S. 37ff. schon beschrieben wurde, geht das Tergum T seitlich in die Apodem- Einstülpung des indirekten Hebers dvmi über und grenzt dann weiter lateral (unter Vermitt- 60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 lung der Sklerite vTS, hTS und RS) an die Co- stalplatte. Verschiedene Gelenke dieses Ter- gum-Seitenbereichs (der sich kaudad weiter bis zum TZ erstreckt) spielen bei der Schubbewe- gung von T eine Rolle: 1) ein Resilingelenk zwi- schen der Tergalbrücke und dem Sklerit vTS (s. Abb. 1a, ohne nahere Bezeichnung), 2) der Gelenkspalt zwischen vTS und hTS, 3) der Gelenkeinschnitt zwischen vTS und RS und 4) eine Membranzone zwischen der medialen Wandung des dvm1-Apodems und T (m, Abb. la) — kaudal geht die Wand des Apodems allerdings sklerotisiert in den Tergalzapfen uber, so daß der untere Teil der Medialwand des dvm1-Apodems zusammen mit dem TZ bewegt: wird und dabei durch die Membran m gegen- uber T einen Bewegungsspielraum erhalt (dies erscheint notwendig, wenn man die oben er- wahnte Torsion des TZ bedenkt; s. dazu auch unten). Da die Zugwirkung von kaudal auf den hTS (t2-Bewegung) starker ist als die Schubwir- kung von proximal auf den vIS (Tb-Bewe- gung), wird der Winkel zwischen vTS und hTS beim Flügelvorschwingen stumpfer — der Ter- gum-Seitenbereich wird also gestreckt. Die Ter- galbrücken-Bewegung wirkt sich dadurch auch nach seitlich auf den Randsklerit aus, bleibt aber anscheinend ohne mechanische Auswirkung auf die CP* selbst, da die (geringe) Bewegung des vTS nach lateral weitgehend in den Gelenken des RS abgefangen wird (der RS kann zwar im Gelenk tl eine Bewegung nach lateral durchführen, sein distal in den Flügel reichen- der Fortsatz — an dem der vca ansetzt — gleitet dabei aber lediglich, in einer Führung der Un- terseite der phCP, an der CP entlang). Beim Flügel-Vorschwingen wird also nur die dhCP (der distale Teil der CP) zusammen mit der RAP und dem Flügel bewegt, der Rest der CP stellt das Widerlager dar und bleibt in Ruhe. Im Verlauf des Vorschwingens wird der proxi- male Membranspalt zwischen CP und RAP ver- größert, der vor dem Gelenk c2 (zwischen der dhCP und CP*) befindliche Gelenkspalt da- gegen verengt (vgl. Abb. 1a > b; 9e > fi). Schließlich schlägt der proximale Rand der dhCP an der phCP an und begrenzt den Bewe- gungsspielraum. Kräfte. — Der dorsale Längsmuskel dlm ist der einzige Muskel (im Mesothorax), der den Flügel um die Drehachse C2/C4 nach vorn zu schwingen vermag (Abb. 1; 9e, f). Der Mecha- nismus ist jedoch (wie schon beschrieben wur- de) nur ın einem kurzen, unteren Abschnitt des Flügelschlags (Ende Abschlag, Beginn Auf- schlag) “freigegeben”; da dem dlm am Anfang der Aufschlagsphase ein erheblich kräftigerer Muskel (der dvm1, s. unten) antagonistisch ge- genübersteht, läßt sich der Zeitpunkt der dlm- Kontraktion auf das Abschlagsende einengen. Das dorsale Apodem des indirekten Hebers dvmi ist zwar einerseits (bedingt durch Ge- lenkspalte und ein Membrangebiet, s. weiter oben) gegenüber der Tergalregion T und gegen- über der CP* beweglich, andererseits jedoch kaudal fest mit dem Tergalzapfen TZ verbun- den. Das dvm1-Apodem wird daher, wenn der Flügel am Ende des Abschlags nach vorn schwingt, aufgrund der Bewegung des TZ kau- dal angehoben und damit insgesamt schräg ge- stellt: dabei wird der dvml, v.a. im kaudalen Bereich, gedehnt. Setzt nun der Flügelaufschlag ein, so wirkt der dvm1 so lange zusätzlich als Rückschwingmuskel des Flügels, bis das Apo- dem wieder seine ursprüngliche Ausrichtung er- reicht hat. Dies ist spätestens dann der Fall, wenn die zwischen phCP und dhCP befindliche Membran des c2-Gelenks straff gespannt ist und kein weiteres Rückschwingen mehr zuläßt. Von da an verteilt der dvm1 seine Kraft (nur noch flügelhebend) gleichmäßig auf die CP und RAP. Der dvmi (zumindest sein kaudaler Ab- schnitt!)) ist somit am Aufschlagsbeginn als An- tagonist des dim flügelrückschwingend wirksam — ein Vorschwingen ist während des Flügel- aufschlags nicht möglich. (Im Falle einer toni- schen dlm-Kontraktion müßte ein Teil der dvmi-Kraft zu Beginn des Aufschlags gegen den dim aufgebracht werden und ginge damit dem Aufschlag verloren; vgl. dazu auch S. 44, 100f.) Ein effizientes Flügelvorschwingen (und voll- ständiges Ausnützen des Vorschwing-Bewe- gungsspielraums ım unteren Abschnitt der Abschlagsphase) erscheint v.a. dann möglich, wenn schon vor der Kontraktion des dlm Kräfte wirksam werden, die den Flügel supinatorisch bis zum Anschlag drehen und dann weiter auch am supinatorischen Anschlag halten (vgl. die “erste Voraussetzung”, S. 58; s. auch Abb. 26e: Anströmung der Flügelunterseite während 1) Eine morphologische Unterteilung des dvml in ei- ne vordere und hintere Portion (die diesem funk- tionellen Unterschied entspricht) konnte nicht ge- funden werden: Der Muskel ist äußerlich einheit- lich — die dorsale Tracheen-Einmündung teilt das Apodem und den Muskel nur scheinbar (vgl. auch S. 114f., Anmerkung 9). Prau: Flugapparat der Libellen 61 des Vorschwingens!). Als ein in dieser Weise nützlicher “Hilfsmuskel” des dim könnte etwa der vca eingesetzt werden, der als Supinator des unteren Schlagumkehrpunktes interpretiert wurde (vgl. S. 51). Der Muskel ist aber wahr- scheinlich noch in einer weiteren Hinsicht für eine Unterstützung und Steigerung der dlm- Funktion geeignet: Da er einen Hebelarm zur Auf-Abschlagsachse P1/P2 besitzt, bremst er den Flügelabschlag gleichzeitig; dadurch kann die Vorschwingbewegung ihrerseits stärker zur Auswirkung kommen (vgl. Abb. 28a, b; zur möglichen aerodynamischen Bedeutung der Vorschwingbewegung s. S. 100). \ Da die Achse T1/T1 in einer senkrecht zur Körperlängsachse stehenden Vertikalebene fest- liegt (d.h.: die Gelenke t1 sind weder nach vorn noch nach hinten relativ zueinander versetzbar), wird der Schub der Tergalbrücke stets auf die Radioanalplatten beider Körperseiten aufgeteilt; beide dlm wirken jeweils sowohl auf den rech- ten als auch linken Flügel. Es ist somit gleichgültig, ob nur ein Muskel sich stark kon- trahiert oder ob beide Muskeln halb so stark ar- beiten — die Muskeln der beiden Körperseiten sind als vollkommen funktionsgleich anzuse- hen. Ihre Auswirkung am Flügel wird jedoch von anderen Kräften mitbestimmt und kann demnach unilateral beeinflußt werden (z.B. durch den vca, s. oben; außerdem — bei Zygo- pteren und Epiophlebia — durch den pa, s. un- ten). Schlagasymmetrien, die zu einer Kippung der Tb um die Körperlängsachse führen, könn- ten außerdem eine (geringe) Rolle spielen (vgl. auch S. 46). Schließlich ist noch eine direkte Beeinflussung der mesothorakalen dim durch das metathorakale Tergum zu bedenken, da die beiden Muskeln kaudal-lateral von der Vorder- randleiste (Antecosta) des Metathorax (acy; Abb. 1) entspringen und der Tergum-Vorder- rand des Metathorax beim Flug mit dem Schlag der Hinterflügel auf- und abbewegt wird (dies wurde in der Abb. 1c angedeutet). Da die dlm jedoch kaudad auseinanderspreizen (sie greifen geradezu “so weit wie möglich” lateral an seitli- chen Vorsprüngen der Antecosta an!)), dürften sich Phasenunterschiede der Vorder- und Hin- terfligel nur geringfügig (über Dehnung oder Stauchung der Muskeln) direkt auf die Vor- schwingbewegung auswirken. Eine starkere in- ') Bemerkenswert ist, daß die metathorakalen dlm der Zygoptera und Anisozygoptera “normal” (ple- siomorph) ausgerichtet sind: bei ihnen besteht die Gefahr einer störenden Einwirkung durch ein fol- gendes Segment ja nicht! direkte Beeinflussung über das Postnotum er- scheint mir dagegen möglich. Nur die Zygoptera und die Anisozygoptera (Gattung Epiophlebia) besitzen im Mesothorax (vom Muskel dvm1, der den Flügel zu Beginn des Aufschlags auf jeden Fall in die Grund- schlagbahnebene zurückschwingt, abgesehen) einen direkten Antagonisten des dim, den Pleu- roalarmuskel pa (Abb. 2; 9e, f)?). Dieser Muskel könnte z.B. für eine unilaterale Einstellung der dlm-Kraftwirkung wesentlich sein. Er könnte darüber hinaus (v.a. wenn der dim nicht kontra- hiert wird) auch für die Erzeugung besonders steiler Flügelschläge eingesetzt werden und würde den Schlagbahn-Spielraum zu besonders großen Winkeln hin erweitern (Abb. 28c). Dar- auf (und auf die mögliche aerodynamische Be- deutung des dlm und pa) soll in der Diskussion (S. 99ff. und S. 109f.) noch eingegangen wer- den. Während der Vor- und Zurückschwingme- chanismus im Hinterflügel-Segment der An- isopteren reduziert ist (s. weiter unten), findet er sich bei Zygopteren und Anisozygopteren in beiden Flügelsegmenten. Der metathorakale Mechanismus dieser Gruppen weist jedoch große Unterschiede (in der tergalen Mechanik und in den Muskelfunktionen) gegenüber dem mesothorakalen auf: Das Hebelapodem HA ist im Metatergum von der davor liegenden Ter- galbrücke (deren Vorderrand die Antecosta III bildet) gelenkig abgesetzt und zur tergalen Mit- tenregion hin abgerückt; das Tergum selbst ist in zwei nach außen vorgewölbte, gegeneinander bewegliche Hälften geteilt, die seitlich jeweils ein Gelenk zum Tergalsklerit (vTS) besitzen. Kontrahiert sich der dlm, so werden diese bei- den Tergalkuppeln auseinander bewegt und die Tergalzapfen TZ dadurch nach innen ge- schwenkt. Der Flügel wird in diesem Fall durch den dim nach kaudal bewegt, also zurückge- schwungen. Der auch im Metathorax antagonis- tische pa bewirkt hier ein Vorschwingen. D.h.: im Metathorax haben die Muskeln dim und pa jeweils die genau entgegengesetzte Funktion wie im Mesothorax?). Vorderer Tergalsklerit 2) Zu diesem Muskel s. auch Anmerkung 14, S. 115. 3) Aus der entgegengesetzten Funktion der serial ho- mologen Muskeln im Meso- und Metathorax kann man schließen, daß sie ursprünglich eine andere (im Falle der dlm wahrscheinlich in beiden Segmenten gleichartige) Funktion besaßen. Dieser Gedanke erscheint mir für die Rekonstruktion des ursprüng- lichen Flugapparates der Pterygoten sehr wesent- lich (vgl. S. 78ff.). 62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 und Randsklerit (der letztere ermöglicht wie im Mesothorax durch sein proximales Gelenk zum vTS die vca-Funktion; vgl. S. 51) werden im Metathorax (ahnlich wie im Mesothorax) von der tergalen Vor-Zurückschwing-Mechanik nicht beeinflußt. Die Tergalbrücke des Meta- thorax (das Ursprungsgebiet der mesothoraka- len dim!) ist andererseits durch ihr kaudales Ge- lenk vom übrigen Metatergum unabhängig und bleibt bei der Vor-Zurückschwingbewegung des Hinterflügels in Ruhe (nicht jedoch bei der Schlagbewegung, bei der die Antecosta III auf- und abbewegt wird; vgl. oben). Die Vor-Zurückschwing-Beweglichkeit der Flügel ist bei Zygopteren und Anisozygopteren in beiden Flügelsegmenten größer als bei An- isopteren. Außerdem sind in beiden Segmenten (vom dvmi abgesehen) jeweils zwei Muskeln (dim und pa) vorhanden. Der Gelenkspielraum im Gelenk c2/c4 ist jedoch im Hinterflügel klei- ner als im Vorderflügel, was darauf hindeutet, daß beim Hinterflügel Schlagbahnveränderun- gen in geringerem Maße möglich sind als beim Vorderflügel. Außerdem läßt sich aus den Größenverhältnissen der Muskeln ableiten, daß die Vorschwingfunktion beim Vorderflügel (dlm) besser entwickelt ist als beim Hinterflügel (pa), während die Rückschwingmöglichkeit des Vorderflügels (pa) geringer ist als diejenige des Hinterflügels (dlm). Hier zeigt sich eine, bei den Anisoptera anscheinend weiter vorangetriebene, verschiedene Spezialisierung der beiden Seg- mente (vgl. auch S. 101 und S. 109f.). Im Mesothorax der Anisoptera fehlt der di- rekte Antagonist des dim, der pa; damit ist der (aktiv einstellbare) Bewegungsspielraum der Flügel (gegenüber Zygopteren und Anisozy- gopteren, die in dieser Hinsicht als plesiomorph anzusehen sind — vgl. dazu auch Anm. 14, S. 115, sowie S. 109f.) sekundär verkleinert. Im Metathorax ist der Vor-Zurückschwingmecha- nismus weitgehend reduziert — der pa fehlt, der dim ist nur sehr schwach entwickelt (rudimen- tar), die Bewegungsmöglichkeit im c2/c4-Ge- lenk ist stark eingeschränkt. In diesem Segment ist außerdem das Metatergum erheblich verein- facht: die Beweglichkeit der tergalen “Kuppeln” fehlt, das verkleinerte Hebelapodem sitzt fest an der Tergalbrücke, vTS und hTS — ursprünglich wohl beide auch im Metathorax vorhanden und durch ein Gelenk getrennt — sind zu einem Sklerit verschmolzen (vgl. auch S. 109f.). Durch diese Veränderungen, die eine direktere Über- tragung der Heber- und Senkerkräfte auf den Flügel mit sich bringen, wird der Metathorax zu einem nahezu reinen Antriebssegment. Die großen Unterschiede im Vor-Zurtick- schwingmechanismus zwischen den Haupt- gruppen der Odonata konnten hier nur ange- deutet werden — sie bedürfen weiterer, verglei- chender Untersuchungen. 2. FLUGELMECHANOREZEPTOREN MECHANISCHE BEANSPRUCHUNG DER REZEPTOREN Das Chordotonalorgan in der Radioanalplatte Auf der Flügelunterseite, wenig distal vom Epifulcrum, befindet sich eine kleine, apodem- artige Einstülpung, der ventrale Ansatz des Flügel-Chordotonalorgans (CH, Abb. 3, 10 und 13a). Von dieser Stelle ausgehend zieht der . Rezeptor nach dorsal-medial zur Kutikula der Oberseite der RAP (dorsaler Ansatz s. Abb. 1a, 11a und 15). Die ventrale Ansatzstelle befindet sich innerhalb einer brückenartigen Sklerotisie- rung, die sich von der Basis des Costalsektors aus nach kaudal bis zur Basis des Cubitalsektors erstreckt und nach proximal und distal durch Membran sowohl vom Epifulcrum als auch vom distalen Flügel abgesetzt ist. Der CH-Ansatz unterteilt diese Skleritbrücke in zwei Teilstücke — ein kürzeres vor der Ansatzstelle und ein län- geres dahinter — die als “Hebelsklerite” des CH aufgefaßt werden (vCH, hCH: vorderer und hinterer Hebelsklerit des Chordotonalor- gans). Das Chordotonalorgan von Anax imperator Leach (der Art, an der auch die elektrophysiolo- gischen Experimente durchgeführt wurden) ist etwa 0,5 mm lang. Phasenkontrast- und poları- sationsoptische Untersuchungen zeigten, daß das Organ bei Anax ungefähr 50 Scolopidien enthält!). Die Zahl der Sinneszellen ıst jedoch höher: eine elektronenmikroskopische Analyse (Risler in Vorb.) ergab, daf$ einige Scolopidien mehrere Sinneszellen enthalten. Da der Rezeptor innerhalb der RAP (distal von der Auf-Abschlagsachse P1/P2) liegt, wird er durch die Schlagbewegungen nicht beeinflußt (jedenfalls nicht direkt; vgl. dazu S. 96f.). Auch bei der Vor-Zurückschwingbewegung wird die RAP als Ganzes bewegt und das CH daher weder gedehnt noch gestaucht. Die Drehungen !) Erhardt (1916) gibt für das CH im Flügel von Coenagrion puella (L.) dagegen nur 16 Stifte an. Prau: Flugapparat der Libellen 63 b Abb. 11. (a) Blick auf die Flügelbasis des rechten Vorderflügels von Anax imperator. Dorsales Ansatzgebiet des Chordotonalorgans punktiert. Das Sensillenfeld CF2 verläuft in der proximalen Fortsetzung der Radius+ Me- dia-Ader in einer Furche (D), das Feld CF1 (®) verschwindet basal hinter der Radius-Kante. Vgl. mit Abb. la, 3 und 15. Maßstab 1 mm; (b) Ansicht der beiden Felder campaniformer Sensillen von vorn-oben; rechter Flügel abgeschlagen (Flügelspitze also links unten). CF1 (®) und CF2 (D) sind fast in ganzer Ausdehnung zu erkennen. Maßstab 0,1 mm. des Flügels um die Längsachse führen dagegen zu deutlichen Längenänderungen des Organs. Dies soll im folgenden für die pronatorischen und supinatorischen Drehbewegungen inner- halb der Schlagphasen und an den Schlagwende- punkten näher untersucht werden. Supiniert der Flügel in der Aufschlagsphase, so wird der Cubitalsektor ventral gegenüber dem Epifulcrum bewegt (vgl. S. 53f.); der nach vorn über die Epifulcrum/Cubitalsektor-Dreh- achse F2 hinausragende hintere Hebelsklerit (hCH) setzt die Skleritbrücke des CH dabei un- ter Spannung und “faltet” sie gewissermaßen in das Lumen der RAP hinein (Abb. 14c—e, 17a)!). Da sich der ventrale Ansatzpunkt des CH dabei sehr genau in Richtung des CH-Ver- laufs nach dorsal-medial bewegt, wird das CH entspannt (bzw. gestaucht). Eine entgegenge- setzte, pronatorische Bewegung des Cubitalsek- 1) Die im Ausgangszustand (0°-Anstellung) schon vorhandene Biegung des Sklerits vCH+hCH nach medial sichert — ebenso wie der ventrad umge- schlagene proximale Rand des Sklerits (Abb. 17a) — die Skleritbrücke gegen eine “Faltung” in die entgegengesetzte Richtung (nach außen). tors führt dagegen zur Dehnung des Organs; dies geschieht z.B. bei verstärkter Kontraktion des Fulcroalarmuskels in der Aufschlagsphase (vgl. S. 54f.). Am oberen Schlagwendepunkt, wenn der Cubitalsektor seinen Anschlag am Arculus erreicht (s.S. 54), ist das CH maximal gedehnt (Abb. 14e—c). È Kontrahiert sich an der Auf-Abschlagswende der hintere Coxoalarmuskel hca, wird der Flügel pronatorisch im Abschlagsdrehbereich bewegt und verwunden (vgl. S. 47tt.). Jetzt be- wegt sich der Costalsektor relativ zum Epiful- crum (um die Epifulcrum/Costalsektor-Dreh- achse E1) und setzt die Skleritbrücke des CH von vorn her unter Spannung; die Ansatzstelle des CH wird erneut nach oben-innen bewegt und das CH entdehnt (Abb. 14c>a; 17a). In der folgenden Abschlagsphase wird die Verwin- dung des Flügels und damit der Dehnungszu- stand des CH wie in der Aufschlagsphase vom Krafteverhaltnis der passiv verwindenden Luft und der (in diesem Fall supinatorischen) Mus- keln bestimmt (vgl. S. 50f.). Die erreichbare maximale Verwindung ist im Abschlagsdrehbe- reich) relativ geringfügig (vgl. S. 54, 102); da der vordere Hebelsklerit außerdem kürzer ist 64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 als der hintere, wird das CH in der Abschlags- phase weniger weit entdehnt als in der Auf- schlagsphase (vgl. Abb. 17b). Am unteren Schlagwendepunkt ist der Rezeptor — bei Er- reichen des supinatorischen Anschlags des Abschlagsdrehbereichs — erneut kurzzeitig maximal gespannt (Abb. 14a—c). Das Flügelchordotonalorgan durchläuft dem- nach an beiden Schlagumkehrpunkten ein Deh- nungsmaximum. Entdehnungen finden in der Auf- und Abschlagsphase statt (Dehnungsmini- ma bei extremer Supination bzw. extremer Pro- nation). Je nach deren Ausmaß ist der Abstand zum Längenmaximum (das an den Wendepunk- ten wohl stets erreicht bzw. durchlaufen wird) unterschiedlich groß. Je nach der Geschwindig- keit der Schlagwendepunktsdrehungen (abhän- gig v.a. von der Kontraktion der Wendepunkts- muskeln; vgl. dazu auch S. 97f.) wird die maxi- male Rezeptor-Länge verschieden rasch erreicht. Felder von campaniformen Sensillen und Sinnesborsten Auf der Oberseite der Radioanalplatte liegen im vorderen Bereich zwei lange Reihen von campaniformen Sensillen (CF1 und CF2; Abb. la, 3, 11 und 12); sıe erstrecken sich etwa in der Flügellängsrichtung und folgen dabei dem Ver- lauf von Radius und Media, die in der RAP noch ein Stück weit zu erkennen sind. Bei Anax imperator Leach sind die Einzelsensillen im ba- salen Abschnitt der Felder äußerlich als bis etwa 20 um lange, stark schlitzförmige Einsen- kungen zu erkennen; nach distal werden die Schlitze kürzer (bis weniger als ein Drittel der Länge der proximalen Sensillen) und z.T. auch oval bis rundlich. Erhardt (1916) beschrieb verschiedene sol- cher “Porenfelder” in der Flügelbasis von Coenagrion puella (L.) und bei anderen Libel- len. Ihr fiel ebenfalls die große Mannigfaltigkeit in der Ausbildung (Form und Größe) des äuße- ren Kutikularapparates auf (runde, ovale oder spaltförmige Gruben). Extrem schlitzförmige Poren bei Hemianax papuensis (Burmeister) be- zeichnete Simmons (1978) als “crevice organs” und verglich sie mit den Spaltsinnesorganen der Spinnen. (Derartig schmale Sensillengruben sind auch bei Anax ganz proximal vorhanden; sie er- lauben wahrscheinlich genauere Rückschlüsse auf die mechanische Beanspruchung und Ver- formung der Kutikula — s. weiter unten.) Die- ser Autor unterschied bei Hemianax papuensis vier Sensillenfelder, alle auf der Flügel-Dorsal- seite, an der Basis der Radius + Media-Ader. Bei den hier untersuchten Arten (Anax impera- tor Leach, Aeshna cyanea Müll.) konnte eine solche Unterteilung jedoch nicht aufgefunden werden — auch nicht bei einem Vertreter der Gattung Hemianax (H. ephippiger Burm.), der zum Vergleich herangezogen wurde. Es fanden sich stets nur zwei Felder, deren Einzelsensillen — ohne größere Unterbrechungen — in ge- schlossener Reihe angeordnet sind: CF2 umfaßt dabei Simmons’ “field 1 + 3 + 4”, CF1 ent- spricht dem “field 2”. Sowohl im Vorder- als auch im Hinterflügel von Anax imperator wur- den je 90 bis 105 Sensillen pro Feld gezählt, wo- bei das CF1 6 bis 12 Rezeptoren weniger als das CF2 aufwies; bei Aeshna cyanea enthielten bei- de Felder (in beiden Flügeln) jeweils etwa 80 Einzelsensillen. Eine genauere raster-elektro- nenmikroskopische Untersuchung zeigte, daß sich die Ausrichtung der Langsachsen der ein- zelnen Sensillengruben im Verlauf der Felder in charakteristischer Weise verändert (teilweise zu erkennen in der Abb. 12). Beide Sensillenfelder werden anscheinend nur durch die Drehbewegungen des Flügels um die Längsachse mechanisch beansprucht!). Die Spannungsänderungen in der Kutikula der dor- salen RAP sind allerdings äußerlich nicht sicht- bar, wie etwa die Langenanderungen des Chor- dotonalorgans (die an der Ein-Auswartsbewe- gung des ventralen Ansatzgebietes direkt beobachtet werden können). Innerhalb der Sen- sillenreihen wurden daher feine Längs-Ein- schnitte in der Kutikula angebracht, die bei gleichzeitiger Drehung des Flügels beobachtet wurden. Ein Aufklaffen der Schnitte im Verlauf der Drehbewegung wurde als Zugbeanspru- chung der Kutikularapparate der dort liegenden Sensillen (quer zum Flügel, in der Längsrich- tung der Gruben) interpretiert. Eine Pronation des Flügels im Abschlagsdrehbereich (welche proximal, wie beschrieben wurde, mit einer Re- lativbewegung der Costalsektor-Basis gegen- über der restlichen RAP verbunden ist), führt danach zu einer Zugbeanspruchung der Kutiku- la quer zum Feld CF1 (Abb. 14c>a); eine Su- pination im Aufschlagsdrehbereich (Bewegung des Cubitalsektors relativ zur restlichen RAP) verursacht dagegen eine Zugbeanspruchung 1) Auch Simmons (1978) vermutete (unter Bezug auf Neville, 1960), daß die Rezeptoren in einem funk- tionellen Zusammenhang mit der Verwindung des Flügels stehen; er gab jedoch keine näheren Erlau- terungen dazu. Prau: Flugapparat der Libellen 65 Abb. 12. (a) Proximaler Abschnitt des Sensillenfeldes CF1 (Ansicht wie in Abb. 11b); hier befinden sich die längsten Kutikular-Schlitze (bis ca. 20 wm). Maßstab 10 um; (b) Mittlere-distale CF1-Sensillen (Ansicht wie in Abb. 11b) — die Kutikulargruben stehen in der Mitte des Feldes schräg zur Radius-Ader, an beiden En- den fast senkrecht zu ihr. Im Hintergrund das Feld CF2. Maßstab 0,1 mm; (c) Aufsicht auf die Basis des Feldes CF2 (Flügelspitze unten). Maßstab 10 um; (d) Aufsicht auf mittlere-distale Sensillen des Feldes CF2. Die Ku- tikulargruben der mittleren Sensillen sind (gegenüber den basisnahen und distalen Gruben) mehr in Flügellängs- richtung ausgerichtet (ähnlich wie beim CF1). CF1 rechts zu erkennen. (Zum gebogenen Gesamtverlauf des Fel- des vgl. auch Pfau, 1983, l.c. Abb. 2—3c.) Maßstab 0,1 mm. | ae ), ist nur andeutungsweise zu erkennen; am unteren Ende wird es von dem borstentragenden Auswuchs BF2 überragt. E kennzeichnet die ungefähre Lage des ventralen CH-Ansatzpunktes. BF1 rechts unten. Vgl. mit Abb. 7 und 17a. Maßstab 0,1 mm; (b) Blick auf das Borstenfeld BF1 von kaudal-ventral (rechter Vorderflügel); BF2 am linken unteren Bildrand noch erkennbar. Maßstab 0,1 mm. Abb. 15 dargestellt). In beiden Drehbereichen nehmen die Zugspannungen bei Drehung zur 0°-Anstellung zurück wieder ab (Abb. 14a—c, ec). Eine kleinere Anhäufung von kurzborstigen Haarsensillen findet sich ganz proximal auf der Unterseite der Costalsektor-Basis (BF1; Abb. 7, 13 und 17a)!). Sie liegt auf einem nach kaudal über das Costalsektor/Epifulcrum-Gelenk el hinausragenden Auswuchs, dem verdickten An- fang der Subcosta. Ahnliche Sinneshaare sitzen “gegenüber” (auf der anderen Seite des Epiful- 3) Wenig distal vom Feld BF1 liegen zwei kleinere Felder campaniformer Sensillen, die hier nicht wei- ter berücksichtigt wurden (cf, Abb. 17a). crum — am proximalen, vorderen Rand der Cu- bitalsektor-Basis) auf dem frontad über das Cu- bitalsektor/Epifulcrum-Gelenk e2 hinausragen- den (kleineren) Fortsatz BF2. Beide Kutikular- auswüchse stellen weiche Stoppstellen (“Anschläge”) für die Verwindungsbewegungen der Flügelsektoren dar. Die inneren Borsten des Feldes BF1 kommen bei starker pronatorischer Verwindung im Abschlagsdrehbereich, diejeni- gen des Feldes BF2 dagegen bei starker Supina- tion ım Aufschlagsdrehbereich, mit der Außen- wand des Epifulcrum in Kontakt und werden abgebogen (Abb. 13b und 14a, e). Selbst bei die- sen extremen Anstellungen wird das Epifulcrum aber offensichtlich nur von wenigen Haarbor- sten direkt berührt, so daß für die Erregung der Abb. 14. Schematische Darstellung der Beanspruchung von Mechanorezeptoren der Radioanalplatte bei Prona- uon und Supination. Blick von distal-dorsal auf die durch einen Querschnitt geöffnete Basis des linken Flügels (der Schnitt verläuft im vorderen Bereich schräg, der Querader cr, folgend). Fulcrum und Epifulcrum wurden eingezeichnet, obwohl sie (bei dieser Ansicht) eigentlich verdeckt sind; die ventrale Verbindung von CP und RAP (mCP-Fortsatz und Gelenk c4) wurde weggelassen (vgl. andere Abb.). Epifulcrum mit Orientierungslinie. © “benutzte” Gelenke, @ “stilliegende” Gelenke; (c) 0°-Anstellung (vgl. Abb. 6); c — a Pronation (a > c Supination) im Abschlagsdrehbereich; c > e Supination (e > c Pronation) im Aufschlagsdrehbereich; a — € Supination des unteren Schlagwendepunkts; e — a Pronation des oberen Schlagwendepunkts. Helle Pfeile kennzeichnen die Bewegungen der Skelettelemente, schwarze die Bewegungen des CH-Ansatz-Stiftes bzw. Zugbeanspruchungen in der Kutikula (zur Übertragung der Kutikularspannungen auf das Sensillenfeld CF2 vgl. auch die Abb. 15). Prau: Flugapparat der Libellen NOILVNOHd NOILVNIdNS 67 68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 ubrigen Haarsensillen die beim Anpressen am Epifulcrum stattfindende Verformung der Vorsprünge eine Rolle spielen könnte. ELEKTROPHYSIOLOGIE DER REZEPTOREN Material und Methode Als Untersuchungstier wurde Anax impera- tor Leach gewählt, die größte einheimische Li- belle, die bei uns im Juli und August an größe- ren Seen stellenweise noch häufig auftritt. Bei kleineren Arten, selbst bei Arten der Gattung Aeshna (die z.T. bis spät in den Herbst hinein zu fangen sind), erwies sich die Präparation als erheblich schwieriger. Da frisch geschlüpfte Li- kMB fa Abb. 15. Schema der Verformungen der kaudalen Radioanalplatte im Aufschlagsdrehbereich und der möglichen Übertragung von Kutikularspannungen auf das Sensillenfeld CF2. Blick auf die RAP eines rechten Flügels von Aeshna. Das kaudale Ansatzge- biet des Fulcroalarmuskels (fa) ist über eine Zone be- sonders verstärkter Kutikula (z) an einem Langsfalz (f) der RAP “aufgehängt” (vgl. S. 54). Das dorsale Ansatzgebiet des Chordotonalorgans (CH) wird ‘durch diesen Falz vermutlich gegen die pronatori- schen (P) und supinatorischen (S) Bewegungen des fa- Ansatzgebietes “abgeschirmt”; weiter proximal wer- den Zug- bzw. Druckspannungen (angedeutet durch die schwarzen Pfeile) möglicherweise über die kauda- le Media-Basis (kMB) auf die CF2-Sensillen übertra- gen. bellen für die Versuche nicht geeignet sind (sie sind weniger robust und fliegen auch schlecht), mußten adulte Exemplare jeweils kurz vor den Experimenten gefangen werden. Dabei handelte es sıch meist um Männchen, seltener wurden auch Weibchen verwandt (Weibchen sind nur in geringer Zahl zu finden, haben aber den Vor- zug, daß sie auch noch abends oder bei schlech- terer Witterung fliegen). Von allen vier Flügeln wurde nacheinander abgeleitet — Vorder- und Hinterflügel zeigten dabei keine wesentlichen Unterschiede. Libellen verhalten sich vor dem Windkanal sehr “launisch” und sind kaum zum Dauerflug zu bringen. Daher wurde zunächst nur vom nicht-schlagenden Flügel abgeleitet. Die Tiere wurden mittels Paraffin auf einer Unterlage fi- xiert und um etwa 30° (Kopf nach unten) ge- kippt, so daß die Terga des schräggestellten Pte- rothorax horizontal ausgerichtet waren (s. Abb. 16). Der waagerecht gestellte (in Schlagmitte befindliche) Flügel wurde wenig distal vom No- dus in die Klammer einer Drehapparatur einge- legt. Mit Hilfe dieser Vorrichtung konnte der Flügel um seine Längsachse gedreht werden — es wurde also die Bewegung imitiert, die als ein- zige zu einer deutlichen Beanspruchung der Mechanorezeptoren CH, CF1 und CF2 geführt hatte. Jenseits einer Winkelscheibe war die Drehachse mit einem linearen Wendelpotentio- meter zur Registrierung des Reizes verbunden. In die dorsale Kutikula der Radioanalplatte wurde ein Fenster geschnitten und der zwischen Tracheenwänden laufende (von den Tracheen “getragene”) sensorische Flügelnerv freipräpa- riert. Zur Ableitung von Potentialen des CH mußte der zu diesem Organ führende Nerven- ast so nah wie möglich an der dorsalen Anhef- tungsstelle des CH (in der Abb. 16 punktiert) in den Haken einer Silberdrahtelektrode (40 um ©) gelegt werden!). Dieser Nerv enthielt in eini- gen Fällen anscheinend dennoch Axone von (distalen) campanıformen Sensillen der Reihen CF1 oder CF2, die durch Ableitung und auch durch anschließende Präparation nachzuweisen waren (s. dazu auch S. 74). Bei den Ableitun- gen vom Feld CF1 bzw. CF2 lag die Haken- elektrode weiter proximal. Selbst bei Durch- trennung anderer Nervenäste (v.a. des zum CH !) Intrazelluläre Ableitungen sind dagegen problema- tisch, da die Radioanalplatte im Abschlagsdrehbe- reich mitbewegt wird (vgl. S. 47ff.), was eine Ver- schiebung der Elektroden im Organ mit sich brin- gen würde. Prau: Flugapparat der Libellen 69 % Ten. err ee DA tes. Abb. 16. Versuchsaufbau. Anax imperator um 30° gegenüber der Horizontalen (h) gekippt, rechter Vorder- flügel in mittlerer Anstellung (0° = Anstellung zwischen den Drehbereichen) in der Drehklammer. In der her- ausvergrößerten, gefensterten RAP sind die freigelegten, zu den verschiedenen Sinnesorganen (CH; CF1,2; BF1,2) führenden Nerven sichtbar; die RAP ist distal des Fensters durchsichtig gedacht, so daß das CH, mit- samt den ventralen “Hebelelementen”, zu erkennen ist. p,s,P,S vgl. S. 71. führenden Nerven) konnte jedoch in keinem Fall völlig sichergestellt werden, daß in den Ableitungen nur CF1- oder CF2-Sensillen ent- halten waren, da der sensorische Flügelnerv (um Anderungen der Mechanik zu vermeiden) distal von den Feldern nicht durchtrennt wurde. Die proximalen Borstenfelder BF1 und BF2 der Flügelunterseite (vgl. Abb. 7, 13, 14a, e und 17a) wurden nicht untersucht. Sie waren leicht vollständig auszuschalten: die Borsten beider 70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Felder wurden abrasiert und der Nerv des Fel- des BF2 meist zusatzlich gekappt (der vom Feld BF1 kommende Nerv trifft weiter proximal auf den sensorischen Hauptnerv und war schon aus diesem Grund in den Ableitungen nicht enthal- ten). Uber den Haken des Silberdrahts und den dort aufliegenden Nerven wurde zur Isolierung ein mit einer Paraffin-Vaseline-Mischung gefülltes, aus einem feinen Infusionsschlauch gezogenes Hütchen gestülpt (vgl. Mohl, 1979). Der Silberdraht mußte über eine Schleife an der Radius + Media-Doppelader festgewachst wer- den — nur auf diese Weise konnte eine Zerrung des Nerven (durch die Drehbewegung der Radioanalplatte im Abschlagsdrehbereich) ver- mieden werden. Die Potentiale wurden nach ih- rer Verstärkung (Differenzverstärker Grass P16) auf dem Oszillographenschirm dargestellt CuP el und mittels eines Schreibers (Physiopolygraph von Schwarzer) dokumentiert. Außerdem dien- te ein Lautsprecher der akustischen Kontrolle. In einigen Fällen wurde eın elektronisches Fen- ster zur alleinigen Darstellung der größeren Spi- kes eingesetzt. Da die Klammer der Drehapparatur den Flügel nur auf einem schmalen Querschnitt er- greift und mitnimmt (Abb. 16), sind zur Rei- zung der proximal im Flügel liegenden Sin- nesorgane CH, CF1 und CF2 weitaus größere Drehbewegungen erforderlich, als sie natürli- cherweise beim Flug (auf der Höhe des Nodus) stattfinden (deshalb die großen Drehwinkel in den Abb. 18—21). Dennoch wird bei diesen weiten Drehurigen proximal in der Radioanal- platte ein natürliches Maf} anscheinend nicht überschritten; dies zeigt sich bei gleichzeitiger Abb. 17. (a) Unterseite der RAP des aufgeschlagenen linken Vorderflügels von Aeshna cyanea. Das Bild zeigt den schrägen Verlauf der Gelenkachsen E1 und E2; das vertieft liegende Gelenk e1 der Achse E1 ist nicht sichtbar — am Gelenk e2 (nach Mazeraten ge- zeichnet) sind die Teile übertrieben weit getrennt. E1 und E2 treten auf der Dorsalseite des Flügels aus, be- vor sie sich kreuzen (dabei steht E1 steiler > Aus- tritt weiter proximal). Nur ein kurzes Stück des CH ist (am ventralen Ansatzpunkt) zu erkennen, dort, wo ein Teil der ventralen Membran entfernt ist. Die ge- strichelten Doppelpfeile deuten die möglicherweise unterschiedlichen Bewegungsebenen (und das unter- schiedliche Ausmaß der Bewegung) des ventralen CH-Ansatzes im Abschlags- und Aufschlagsdrehbe- reich an (vgl. S. 102). Die Pfeile an den Gelenken der CH-Hebelsklerite kennzeichnen die Schub-Punkte (bei Flügelverwindung) der R+M- bzw. CuP-Ader; beide liegen unterhalb der jeweiligen Epifulcrum-Ge- lenkachse. cf: 2 kleine Felder von campaniformen Sensillen in der Sc-Basis. Membran punktiert; (b) Eine direkte Messung der Lange des CH bei verschiedenen geometrischen Anstellungen des Flügels war noch nicht möglich. Das Schema zeigt eine (qualitative) Re- konstruktion der Umsetzung eines durch beide Dreh- bereiche gehenden äußeren Sinus-Drehreizes (oben) in Längenänderungen des CH (unten). Ausgezogene Linie: künstliche Flügeldrehung; gestrichelte Linie: durch elastische Kräfte modifizierter (naturgetreue- rer) Verlauf; log: Länge des CH; Ab-Dr, Auf-Dr: Abschlags-, Aufschlagsdrehbereich; «a: geometri- sche Flügelanstellung. Vgl. S. 62ff. und S. 102f. Prau: Flugapparat der Libellen 71 Beobachtung der auf der Flügelunterseite lie- genden Gelenke der Radioanalplatte. Eine wesentliche Frage ist, ob die erzeugten Reize sonst als angenähert physiologisch be- trachtet werden können. Einerseits hatte sich ja ergeben, daß die beiden Drehbereiche unter- schiedliche Drehachsen aufweisen, wobei eine der Achsen (und damit die im Raum verlaufen- de “eigentliche Drehachse”) sich zudem noch während der Flügeldrehung in ihrer Ausrich- tung verändert (s.S. 50); andererseits sind zu- mindest die passiven Kräfte kaum genau zu imi- tieren, da sie am ganzen Flügel angreifen und von proximal nach distal mit dem Quadrat der Umfangsgeschwindigkeit anwachsen. In dieser Hinsicht kann die verwendete Drehvorrichtung (mit ihrer konstanten Drehachse und nur schmalen Kontaktstelle der Drehklammer) si- cher keinen naturgetreuen Reiz erbringen. Da künstlich erzeugte Flügelverwindungen aber so- wohl von proximal wie von distal aus zu den auf S. 62ff. beschriebenen Beanspruchungen der Rezeptoren führen (die Ein- und Auswärtsbe- wegungen des CH-Ursprungs sind dabei leicht über einen Spiegel direkt zu beobachten), kann angenommen werden, daß die Unterschiede zu den natürlichen Beanspruchungen v.a. quantita- tiver Natur sind. Die auf der Flügelunterseite befindlichen Scharniergelenke el und e2 sorgen anscheinend dafür, daß Flügelverwindungen proximal stets in prinzipiell gleicher Weise ablaufen. Weite Drehungen des Flügels, durch beide Drehbereiche hindurch, wie sie etwa in Abb. 18a, b dargestellt werden, finden bei fliegenden Libellen an den Schlagwendepunkten statt. Allerdings sind Pronation und Supination beim Flug natürlich jeweils durch eine Schlagphase voneinander getrennt. Pronation und Supina- tion laufen außerdem sicher erheblich schneller ab. Um mit der Klammer der Drehapparatur entsprechend schnelle Drehungen (und Rezep- tor-Reizungen) zu erreichen, müßte der Flügel im Versuch (infolge der oben beschriebenen Untersetzung vom Ort der Drehklammer zur Flügelbasis hin) mit einer Geschwindigkeit von schätzungsweise 10.000 bis 20.000 Grad/Sekun- de gedreht werden. Da dies technisch im Mo- ment kaum zu lösen ist, wurden zunächst nur manuelle, relativ langsame Drehreize gegeben. Ableitungen vom Flügel-Chordotonalorgan Die Ableitungen, bei denen vom Nerv des Chordotonalorgans, dicht beim dorsalen Aus- trittsort aus dem Organ (Abb. 16), abgeleitet wurde, können (abgesehen von den Einzelfäl- len, die auf S. 73 beschrieben werden) als rei- ne CH-Ableitungen betrachtet werden. Sie stel- len, wie auch fast alle anderen Ableitungen, Summenableitungen dar. In den Abb. 18a und b werden manuelle Si- nusdrehungen durch beide Drehbereiche wie- dergegeben. Die Drehrichtungen innerhalb der Drehbereiche sind (in Abb. 18a) in der oben dargestellten Reizspur durch die Buchstaben p und s (Pronation bzw. Supination im Ab- schlagsdrehbereich) und S und P (Supination bzw. Pronation im Aufschlagsdrehbereich) ge- kennzeichnet (vgl. auch Abb. 16). Auf der Win- kelskala am Bildrand, welche die Stellung der Drehklammer auf der Höhe des Nodus wieder- gibt (Abstand zwischen den Teilstrichen 50°), stehen pmi und Sx für extrem pronierte bzw. supinierte Anstellungen im Abschlags- bzw. Aufschlagsdrehbereich. Die Abb. 18a und b zeigen mehr oder weni- ger dichte Spike-Ansammlungen, die zwischen den extremen Flügelanstellungen auftreten. Da der Flügel bei Pronation (p,P) und Supination (S,s) jeweils die 0°-Anstellung durchläuft (wobei die direkte Beobachtung der Flügelunterseite zeigt, daß das ventrale Ursprungsgebiet des CH in der O°-Anstellung maximal aus der Radio- analplatte herausbewegt ist), treten die Impulse offensichtlich da gehäuft auf, wo der Rezeptor stark gedehnt ist (vgl. Abb. 14, 17b und S. 63f.). Der Flügel ist bei 0° (der Anstellung maximaler CH-Dehnung) auf der Höhe des Nodus ungefähr 25° gegenüber der Horizonta- len supiniert; das bedeutet, daß der 0° entspre- chende geometrische Anstellwinkel bei einem horizontal ausgerichteten Tier etwa 55° Supina- tion betragen würde (vgl. S. 68 und Abb. 16). Die CH-“Bursts” sind aus Spikes unter- schiedlicher Höhe zusammengesetzt, stammen also von verschiedenen Einzelsensillen. Das von Reiz zu Reiz sich ändernde äußere Impulsbild deutet darauf hin, daß das Organ die Unregel- mäßigkeiten des manuellen Reizes wiedergibt. Stärker beschleunigte Drehungen (etwa die er- ste Pronation der Abb. 18a) weisen z.B. beson- ders kurze, dichte Impulsansammlungen auf. Bei langsamen Drehungen sind die CH-Spikes dagegen über den ganzen Drehspielraum (SmaxPma, und zurück) verteilt — die Anhäu- fung um 0° herum ist weniger deutlich (Abb. 18b). Bei Treppen-Reizen durch beide Drehbe- reiche gibt das CH auf jeder Reizstufe Potentia- le ab (Abb. 18c); es wird demnach in beiden Drehbereichen sowohl bei Pronation als auch 72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 b 1 sec eere mn 100 Cc 50 O 50 100 sec LUE ty I d + a! ‚1sec |’ ae a [ssf ei Jr de lean Gee mee 1sec isn ee Abb. 18. (a-e) Ableitungen vom Chordotonalorgan von Anax imperator bei manueller Flugeldrehung um die Längsachse. Impulse nachgezeichnet. In e wurde der Beginn der (sehr geringfügigen) Reize durch Pfeile mar- kiert. bei Supination gereizt. In den Exirembereichen (Pmax bzw. Smax auf der Winkelskala) sind die Antworten etwas schwächer oder können auch ganz fehlen. Auffällig ist, daß die Impulse ent- weder über die ganze Länge der einzelnen Reiz- rampen verteilt sind oder nur am Beginn und Ende abgegeben werden (vgl. auch Abb. 19b). Um 0° ıst das CH anscheinend besonders emp- findlich (Abb. 18d, e): Hier feuern Sensillen noch bei besonders geringfügigen und langsa- men Drehungen des Flügels. In der Abb. 18e wurde der Flügel z.B. nur etwa 5° weit gedreht; diese Winkeländerung wird zur Flügelbasis hin noch beträchtlich herabgesetzt und ist dort un- ter dem Binokular nicht mehr als Bewegung des CH-Ansatzes zu erkennen (in diesem Fall war daher nicht zu entscheiden, in welcher Weise Dehnung und Entdehnung des Rezeptors auf- einander folgten). Ableitungen von anderen Rezeptoren, wahrscheinlich campaniformen Sensillen Die folgenden Ableitungen stellen Einzelbe- obachtungen dar und werden vorerst entweder distalen bis mittleren Sensillen des CF2-Feldes oder CF1-Sensillen zugeschrieben. Die Argu- mente dafür werden weiter unten zusammenge- faßt. Prau: Flugapparat der Libellen 73 Bei zwei Ableitungen (von verschiedenen Tieren), bei welchen ebenfalls weit distal, vom Nerv des Chordotonalorgans, abgeleitet wurde (die Hakenelektrode lag etwa wie in der Abb. 16 dargestellt), feuerten bei Supination, in der Nähe von So Sensillen mit einem von den CH-Sensillen abweichenden Zeitverhalten (Abb. 19). Sie waren im Lautsprecher deutlich von den in der selben Ableitung auftretenden, mehr unregelmäßigen (“stotternden”) Impuls- mustern der stark phasischen CH-Sensillen zu unterscheiden: bei Sinusreizen bildeten sie dich- te Bursts mit z.T. sehr regelmäßiger Folge der Einzelimpulse, die mit dem Beginn der Rück- drehung (P) endeten (Abb. 19a, c)!). Treppen- reizungen zeigten, daß diese Sensillen zu Beginn jeder Reizstufe neu erregt wurden und jeweils mit einer hohen Anfangsfrequenz feuerten (Abb. 19b). Die Erregung ging bei längeren In- tervallen zwischen den Reiztreppen oder zwi- schen der S- und P-Drehung relativ schnell auf Null zurück (vgl. Abb. 19b, 2. Ableitung, und 19d). Bei Sinusreizen kam der Erregungsabfall der Einzelsensillen dagegen nıcht zum Aus- druck, so daß die Impulse dort ohne Lücke bis zum Beginn der P-Drehung aneinander an- schlossen. Offensichtlich folgten dabei mehrere, verschiedene Sensillen dicht aufeinander (vgl. Abb. 19c, d). Das unterschiedliche äußere Im- pulsbild dieser Rezeptoren bei den Sinusreizen erklärt sich wohl v.a. aus den unterschiedlichen Dreh-Amplituden und -Geschwindigkeiten. In der Abb. 19d ist z.B. zu erkennen, daß ein ein- zelnes Sensillum in Abhängigkeit von der Dreh- amplitude (und Geschwindigkeit?) verschieden stark erregt wurde. Bei Drehungen höherer Geschwindigkeit zeigten die Ableitungen dich- tere, bei gleicher Drehamplitude zwangsläufig kürzere Bursts (Abb. 19c). Der Beginn der Bursts hängt anscheinend wesentlich vom Aus- gangswinkel der Drehung ab: ging der nächste Drehreiz z.B. von einem größeren geometri- schen Anstellwinkel (näher bei $,,,,) aus, so ver- schob sich der Erregungsbeginn zu einem größeren Winkel hin (auch dann, wenn schon früh eine dem vorigen Reiz entsprechende Drehgeschwindigkeit erreicht wurde; Abb. 19d, e). Diese Einzelbefunde sprechen dafür, daß der Gesamtverlauf des Reizes (d.h. sowohl Anfang und Ende als auch Geschwindigkeit der Dre- !) Nur bei über den Bereich der ableitbaren Sensillen hinausgehenden (oder auch bei unphysiologisch weiten) Drehreizen endeten die Bursts schon vor dem Beginn der P-Drehung. hung) für die Erregung der Sensillen von Bedeu- tung ist (vgl. dazu auch S. 103t.). Von ganz ähnlichen Sensillen (mit entspre- chendem Zeitverhalten) konnte auch bei Prona- tionsdrehungen im Abschlagsdrehbereich, in der Nahe von p,,,,, abgeleitet werden (nicht ab- gebildet). Auch hier feuerten die Sensillen nur bei Drehung in Richtung Anstellextrem und en- deten bei Beginn der Rückbewegung (in diesem Fall dem Beginn der s-Drehung). In einigen Fällen wurde weiter proximal vom sensorischen Flügelnerv abgeleitet; die Haken- elektrode befand sich dabei vor der Aufgabe- lung des Haupt-Nervenstammes in die zu den verschiedenen Rezeptorfeldern führenden Äste (jedoch distal von der Abzweigung des zum Feld BF1 führenden Nerven). Auch hier traten Sensillen mit einem mehr phasisch-tonischen Zeitverhalten auf. Bei gleichmäßiger und lang- samer pronatorischer Drehung durch den Abschlagsdrehbereich steg die Impuls- zahl/Sekunde von 0° an allmählich an, wobei die im Zeitverlauf unterschiedlichen Spikegrößen auf ein sukzessives Ansprechen immer neuer Sensillen hinweisen (Abb. 20a). Stufenreize zeigten, daß bestimmten Flügelanstellwinkeln im Abschlagsdrehbereich bestimmte Summen- erregungszustände zugeordnet sind (Abb. 20b). Bei dieser Ableitung wurde der Flügel jeweils um 30° weiter proniert, dann wurde eine Sekun- de gewartet und aufgezeichnet. Die Erregung blieb auf jeder Stufe über mehrere Sekunden re- latıv konstant; nur bei 60° und 90° sieht man in der zweiten Hälfte der Ableitung einen gerin- gen Erregungsabfall (möglicherweise ein Hin- weis darauf, daß die distalen Sensillen der Felder schneller adaptieren; vgl. auch Abb. 19b, d und weiter unten). Von ähnlichen, “phasisch-toni- schen” Sensillen konnte auch bei supinatori- schen Drehungen im Aufschlagsdrehbereich (S) abgeleitet werden — hier jedoch nicht ab 0°, sondern erst ab 50° (s. Abb. 21 und weiter un- ten). Alle diese vom Erregungsmuster der stark phasischen CH-Sensillen abweichenden Ablei- tungen werden vorerst campaniformen Sensillen zugesprochen. Während die auf Pronation an- sprechenden Sensillen des Abschlagsdrehbe- reichs als proximale bis distale Sensillen des Fel- des CF1 angesehen werden können, stammen die bei Supination im Aufschlagsdrehbereich abgeleiteten Impulse wohl von mittleren bis di- stalen CF2-Sensillen (von distalen Sensillen ın den Fällen, in denen scheinbar allein vom CH- Nerv abgeleitet wurde). Für diese Interpreta- 74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Wt max HA LE er O 1sec HL HNE cai i tion spricht folgendes: 1) Die mechanische Beanspruchung der Sensillengebiete (S. 64ff.) weist darauf hin, daß in der Kutikula Zugspan- nungen in der Langsrichtung der Kutikulargru- ben auftreten — entweder bei Pronation im Abschlagsdrehbereich (CF1) oder bei Supina- tion im Aufschlagsdrehbereich (CF2) —, die von proximal nach distal fortschreiten (vgl. auch Abb. 14 und 15). Diese verursachen wahr- scheinlich — tber Druckspannungen senkrecht zur Grubenlangsachse und Deformationen der Dendriten-Tubularkörper (vgl. z.B. Zill & Mo- ran, 1981) — eine sukzessive Reizung der auf- einanderfolgenden Sensillen. 2) Die “typischen” Ableitungen vom CH stellten stark phasische Antworten dar, die v.a. im mittleren Anstellbe- reich (um 0°), also bei stärker gedehntem Re- zeptor, auftraten; diese Sensillen wurden in bei- den Drehrichtungen erregt. Die Sensillen mit dem abweichenden (phasisch-tonischen) Zeit- verhalten traten dagegen in beiden Drehberei- chen jeweils nur in einer Richtung, namlich zum Anstellextrem hin, auf; in dieser Richtung wird das CH zunehmend entspannt und reagiert im- mer schwacher. 3) In den Fallen, in denen vom CH-Nerv allein abgeleitet wurde und dennoch Sensillen mit phasisch-tonischem Zeitverhalten feuerten (s.S. 73), konnte bei anschließender Präparation gezeigt werden, daß von distalen CF2- (bzw. CF1-) Sensillen stammende Nerven an den Nerv des CH angeschlossen waren. (Das Verzweigungsmuster erwies sich überhaupt als äußerst variabel: die Axone der einzelnen Sen- sillen der campaniformen Sensillenfelder sind keineswegs immer streng gebündelt, sondern nehmen z.T. “Umwege”; so können CF2-Ner- venäste z.B. auch an den BFi-Nerv an- schließen.) Anscheinend wurden gerade diese Sensillen — der Erwartung entsprechend in den extremen Anstellbereichen — erregt. 4) Das se- Prau: Flugapparat der Libellen 75 1 sec abs U ee RM HI a en) d 2 ' max dd 1sec Abb. 19. (a-f) Gemischte Ableitungen vom Chordotonalorgan und (vermutlich) distalen Sensillen des Feldes CF2. In (a) und (c) sind die CF2-Bursts durch Balken gekennzeichnet (in der Reizspur steht in (a) am Winkelort ihres Beginns ein x); in (e) ist der Drehabschnitt mit CF2-Impulsen in der Reizspur verdickt wiedergegeben. Vgl. S. 72f. In (f) sind neben CF2-Sensillen (die durch Punkte unter der Ableitspur gekennzeichnet wurden) verschiedene, bestimmten Winkelabschnitten der Drehung zugeordnete CH-Impulse (f,U) zu sehen (s. dazu S. 102). rielle, dichte Aufeinanderfolgen der Potentiale der Sensillen entspricht ihrer Reihenanordnung. Zeitlich spätere (jeweils bei größeren geometri- schen Anstellwinkeln auftretende) Antworten können anscheinend weiter distal liegenden Sensillen zugeordnet werden (s. oben). Bei weit proximal angelegter Elektrode zeigten von 0° ausgehende Supinationsdrehungen allerdings nie von Beginn an Impulse, d.h., die basalen CF2-Sensillen fehlten immer. Dieser Teil des CF2-Feldes mußte durch die Präparation (im Gegensatz zum Feld CF1) starker beschadigt werden (vgl. Abb. 16)! 5) Andere Rezeptoren, die ebenfalls mit den Flügeldrehungen in einer Beziehung stehen könnten (BF1, BF2; vgl. S. 66ff.), waren ausgeschaltet (s.S. 69f.). 3. EVOLUTION DER FLUGAPPARATE DER ODONATEN, EPHEMEROPTEREN UND NEOPTEREN In diesem Kapitel soll auf einige Aspekte der Evolution der verschiedenen Flugmechanismen der Pterygoten eingegangen werden. Aufgrund 76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 a rn au ta ie | AE Il] 1sec Où LA LL LA LL TL LA LT NU (LL 1 si U I 30 Att AA AAA Att] LL I Po" slee HHH Abb. 20. (a) Allmähliche Pronation im Abschlagsdrehbereich (untere Spur; 0°-Linie gestrichelt mitgeführt) und Ableitung von (diesem Drehbereich zugeordneten) Sensillen ue Feldes CF1 (vgl. S. 73); (b) Ableitung von entsprechenden Sensillen wie in (a) bei unterschiedlichen (statischen) Flügelanstellungen im Abschlagsdrehbe- reich (vgl. S. 73). | 1sec Ss SET, es Loose IPSOA ge TI Oo fn TI È S a A UE ANN CATT DEL OC TEL AT e a mn mn nn Abb. 21. Ableitungen, die mittleren bis distalen Sensillen des Feldes CF2 zugeordnet werden; Supinations- drehungen des Flügels im Aufschlagsdrehbereich (vgl. S. 73ff.). 1) Hennig (1969, l.c. S. 34) führte für Gruppenbe- der fast ganz fehlenden Fossilüberlieferung und EE zeichnungen eine sehr wesentliche Präzisierung der Notwendigkeit, von gut gesicherten mo- k ann fe Dec MEL ACID Re i CE ein: er kennzeichnet diejenigen Gruppen mit einem Done NDR 5 3 Sternchen (z.B. *Trichoptera), die auf den letzten bei v.a. ein Vergleich der SATE Gruppen as gemeinsamen Vorfahren aller rezenten Arten grunde gelegt werden (s. dazu die grundsätzli- zurückgehen und unterscheidet sie von Gruppen chen Überlegungen von Hennig, 1969)!). Da ohne *, die auch die fossilen sog. “Stammgruppen” Prau: Flugapparat der Libellen 77 Locusta } Cetonia Apis Calliphora Abb. 22. Unterschiedliche Pronations-Supinations-Mechanismen bei Neopteren. Flügel in supinierter Anstel- lung gezeichnet. Der dick schraffierte Basisabschnitt bildet bei einer Verwindung des Fliigels jeweils das Wider- lager; beim Flügelschlag werden die thorakalen Antriebskräfte über diesen Teil auf den “Verstellflügel” (weit schraffiert) übertragen. Lage der Drehachse des Flügels gestrichelt angedeutet. Die Pfeile in den Muskeln deuten eine Funktion beim Aufschlag (Pfeilspitze nach oben) oder Abschlag (Pfeilspitze nach unten) an; dabei wird eine zugfederartige Wirkung der wohl tonisch-aktiven Muskeln angenommen (Ausnahme: die phasischen Basalar- und Subalarmuskeln von Locusta und der subl von Cetonia). Pronatoren dunkel (eng punktiert), Supinatoren hell (weit punktiert). Nur der Muskel pt3 (M85) von Locusta wirkt in den beiden Schlagphasen unterschiedlich, beim Abschlag supinierend, beim Aufschlag pronierend, jeweils entgegengesetzt zu der den Flügel passiv dre- henden Luftanströmung (wobei die Drehwirkung beim Abschlag vermutlich noch durch eine hinzukommende “Zugfeder-Wirkung” verstärkt wird). bas Basalarmuskeln; sub Subalarmuskeln; pt3 Pterale-3-Muskeln; pt4 Pterale-4-Muskeln; ts Tergosternalmuskel; die tiefgesetzten Zahlen sollen keine Homologievorstellung ausdrük- ken. Die Muskeln für Schlagbahnveränderungen (weitere Pterale-3-Muskeln und Basalarmuskeln bei Apis und Callıphora) wurden weggelassen. Vgl. Pfau, 1977b, 1978a; Pfau & Honomichl, 1979; Pfau, 1977a und in Vorb. unsere Kenntnis über die Funktionsweise der rezenten Fiugapparate jedoch immer noch sehr lückenhaft ist, ist diese Basis sehr schmal. So muß der Vergleich hier weitgehend auf den Flügelantrieb beschränkt bleiben; eine syntheti- sche Theorie, welche die Flügel-Stellfunktionen mit einbezieht, ist vorerst höchstens in Umris- sen möglich. Dies hängt auch damit zusammen, daf} innerhalb der großen Gruppe der Neoptera sehr unterschiedliche Flügelstellmechanismen verwirklicht sind: So hat sich z.B. gezeigt, daß die Drehbewegungen des Flügels um die Längs- enthalten. In der vorliegenden Arbeit handelt es sich in der Regel um *Gruppen; nur an einigen Stellen, an denen ich ein Mißverständnis auf jeden Fall vermeiden möchte, wird entweder ein * an- gefügt oder ausdrücklich vermerkt, daß Gruppen im weiteren Sinne (d.h. incl. Stammgruppe) ge- meint sind. achse bei Orthopteren, Coleopteren, Hyme- nopteren und Dipteren in ihrer Mechanik größere Unterschiede aufweisen, und daß in diesen Gruppen z.T. nicht-homologe pronatori- sche bzw. supinatorische Muskeln eingesetzt werden (Abb. 22; vgl. auch Pfau, 1977a, b; Pfau & Honomichl, 1979). Entsprechendes trifft für die Muskulatur und Mechanik der Flügelschlagbahnänderungen zu (eigene, nicht veröffentlichte Untersuchungen). Der Grund- plan der an der Basis der Neoptera vorhandenen Stellmechanismen muß also erst noch rekon- struiert werden. Da selbst innerhalb einzelner Ordnungen der Neoptera keine Einheitlichkeit besteht, müßten zunächst für diese Gruppen ge- nauere vergleichende Bearbeitungen und Grundplanrekonstruktionen erfolgen. Die im weiteren vorgenommene Rekonstruk- tion einiger wesentlicher Teile des Ur-Flugap- parates der Insekten, und ihrer Abwandlungen 78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 auf dem Weg zu den rezenten Gruppen, ist ein Versuch, die z.T. großen Lücken zwischen den rezenten Flugapparat-Typen zu schließen. Er muß weitgehend hypothetisch bleiben. Wenn hier dennoch an verschiedenen Stellen bis ın Einzelheiten gegangen wird, so v.a. deshalb, um aufzuzeigen, daß die These eines polyphyleti- schen Ursprungs des Insektenflugs (Matsuda, 1981) keineswegs zwingend ist (sie geht außer- dem von einigen falschen Voraussetzungen aus vgl. dazu auch S. 105ff.). Der ursprüngliche Antriebsmechanismus der Pterygoten Bei den drei rezent existierenden Hauptgrup- pen der Pterygoten, den Odonaten, Epheme- ropteren und Neopteren, sind zwei verschiede- ne Antriebs-Grundprinzipien verwirklicht. Sie sollen hier (schlagwortartig) als “Tergalplatten- Mechanismus” TPM (Odonata) und “Tergal- wölbungs-Mechanismus” TWM (Ephemeropte- ra, Neoptera) bezeichnet werden. Der TPM ist dadurch gekennzeichnet, daß das Tergum beim Flügelschlag als Ganzes auf- und abbewegt wird (Abb. 24b) und die Flügel- basis demzufolge über eine längere Strecke (bzw. über mehrere Gelenkstellen) scharnierar- tig mit dem Tergum artikuliert. Als antagonisti- sche Antriebsmuskeln existieren indirekte He- ber (am Tergum ansetzende Dorsoventralmus- keln, die primar wohl uber die ganze Breite des Schlaggelenks verteilt waren) und direkte, am Flügel ansetzende Senker (“Basalar-” und “Sub- alarmuskeln”). Der TWM führt dagegen über eine Aufwol- bung und Abflachung des Tergum zum Ab- und Aufschlag, wobei der Flügel nur durch ei- nen Teil des Tergalrandes, den hauptsächlich auf- und abbewegten mittleren bis hinteren Be- reich, gehebelt wird (Abb. 23 und 24c, d). Membranöse seitliche Einschnitte im Tergum (Tergalspalte) verbessern die Verwölbbarkeit des Tergum und somit seine Hebelwirkung — sie konzentrieren sie auf einen relativ kurzen Abschnitt der Flügelbasis. Als Hauptmuskeln des TWM fungieren am Tergum ansetzende Dorsoventralmuskeln (indirekte Heber) und dorsale, rein tergale Längsmuskeln (indirekte dvm dim Abb. 23. Das Wirkungsprinzip des Tergalwölbungs-Mechanismus (TWM) kann nur in einer Seitenansicht des Thorax verdeutlicht werden. Tergum und Sternum punktiert, Pleuralleiste schwarz hervorgehoben. Nur die in- direkten Muskeln wurden eingezeichnet. Prau: Flugapparat der Libellen 79 Senker). Außerdem können phasische direkte Senker als Synergisten der dorsalen Längsmus- keln beteiligt sein (mit deutlicher Abschlagswir- kung v.a. dann, wenn sie in ihrer Zugrichtung mit den Dorsoventralmuskeln übereinstimmen, und wenn ihr Zugpunkt am Flügel der tergalen Hebelstelle genahert ist); sie haben jedoch meist zusätzliche Funktionen als Stellmuskeln und sind in abgeleiteten Gruppen (etwa Dipteren) auch zu reinen (tonischen) Stellmuskeln gewor- den. Das hier als TWM bezeichnete Antriebsprin- zip wird ın der Literatur (vgl. etwa Hadorn & Wehner, 1974; Seifert, 1975) häufig falsch dar- gestellt. So wird z.B. die Aufwölbung des Ter- gum durch die dorsalen Längsmuskeln, die ın der Tierlangsrichtung geschehen muß (vgl. Abb. 23 und 24d), in eine Querschnittsebene des Thorax verlegt (s. z.B. Seifert, 1975, Abb. 148); eine zunehmende Quer-Verwölbung des Ter- gum würde aber dem Abschlag (den sie bewir- ken soll) sogar entgegenarbeiten und ist außer- dem kaum durch eine Kontraktion der Längs- muskeln zu bewerkstelligen. Es ist auch nicht korrekt, wenn man den Mechanismus des Flügelschlags der Insekten als “Deckel-Topf- Mechanismus” darstellt und verallgemeinert, da dieses Prinzip eigentlich nur bei den Odonaten (als TPM) verwirklicht ist. V.a. bei stark spezia- lisierten Gruppen, die den Flügel über einen kaudalen “Scutellarhebel” antreiben (Dipteren, Hymenopteren, Ephemeropteren), spielt eine “Deckel-gegen-Topf”-Bewegung für den Flü- gelschlag keine Rolle mehr: die Bewegung der Scutellarhebel wird hier durch eine komplizierte Verformung des Tergum und der Pleuren (d.h. fast des gesamten Thorax!) bewirkt. Da die indirekten Senker (dim) und Heber (dvm) und die direkten Senker (bas, sub) in allen Pterygoten-Hauptgruppen vorhanden und am Flügelantrieb beteiligt sind (im Falle des dlm der Odonata kann auf eine ursprüngliche Ab- schlagsfunktion geschlossen werden; s.S. 61), kann davon ausgegangen werden, daß diese Muskeln auch bei der Ahnform der *Pterygota als Flügelantriebsmuskeln vorhanden waren !). Das wiederum könnte bedeuten, daß primar ein Schlagmechanismus existierte, der sich noch beider Möglichkeiten bediente, also sowohl das 1) Dies steht im Gegensatz zu der verbreiteten Auf- fassung, daß an der Basis der Pterygoten eine über- wiegend direkte Schlagmuskulatur verwirklicht war (vgl. z.B. Pringle, 1957, S. 4; Kaestner, 1972, S. 71). TPM- als auch das TWM-Prinzip nutzte. Die Funktionsfähigkeit eines solchen Ur-Flugappa- rates (“TPM + TWM”, Abb. 24a), der, wie sich im weiteren zeigen soll, eine relativ zwanglose Ableitung der (effizienteren) rezenten Flügelan- triebsmechanismen erlaubt, setzt bestimmte Konstruktionsmerkmale voraus: So muß z.B. sowohl der TPM- als auch der TWM-Antriebs- anteil über ein eigenes Schlaggelenk und eine eigene Schlagachse verfügt haben; in der Abb. 24a wurden daher zwei pleurale Flügelgelenke, a und b, und zwei Schlagachsen (A/B und B) eingezeichnet. Da das Tergum auf der Höhe des (etwa ın der Flügelmitte liegenden) Gelenkes b beim Flug stärker auf- und abbewegt wird als weiter vorn, auf der Höhe von a (s. Pfeile in Abb. 24a), ist zusätzlich die Annahme einer in- nerhalb der Flügelbasis verlaufenden, schrägen Gelenkzone notwendig. Sie trennt einen vorde- ren, basalen Teil des Flügels, den “Basis-Skle- rit” (BAS), vom restlichen Flügel. Vorn sitzt dieser Sklerit dem vorderen pleuralen Gelenk- kopf auf (und bildet das Gelenk a), hinten be- sitzt er — an der Stelle e, dicht beim hinteren Schlaggelenk b — eine zweite gelenkige Kon- taktstelle zum Pleurum. Eine solche Gelenk- und Achsenanordnung erlaubt es, daß beide Antriebsmechanismen (TPM: Bewegung des ganzen Flügels um A/B — TWM: Bewegung des distal von BAS liegenden Flügelabschnittes um B) gemeinsam und auch relativ gleichwertig an der Flügelschlagbewegung beteiligt sein kön- nen (würde z.B. der Sklerit BAS an anderer Stelle artikulieren — etwa bei e’ oder e”, vgl. Abb. 24b und d —, so würde dies entweder den TWM- oder den TPM-Antriebsanteil in seiner Effektivität schwächen; s. dazu auch die folgen- den Kapitel). Die Schlagachse B des TWM darf in einem solchen System nicht als flügelfeste Achse ausgebildet sein: eine feste Achse B wür- de bei der Bewegung des (ganzen) Flügels um die Achse A/B ihre Ausrichtung zur tergalen Hebelstelle verändern und wäre nur in einem kurzen Schlagphasenabschnitt günstig ausge- richtet; eine “Schlagachse mit Bewegungsspiel- raum” weist dagegen diesen Mangel nicht auf. Man kann daher weiter postulieren, daß das Ge- lenk b kugelgelenkartig ausgebildet war, und daß außerdem der distal von BAS liegende Ge- lenkspalt nach vorn (zur Costa hin) breiter wur- de, um dem Flügel — für seine Bewegung ge- genüber dem BAS, um die nicht-flügelfeste Achse B — einen Spielraum zu geben. Der Vor- derrand der Gelenkmembran konnte durch ein auf beiden Seiten mit Gelenken (c und d) verse- 80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 henes Teilstiick der Costa, die Humeralplatte HP, eine gewisse Festigkeit behalten und gleichzeitig die für die Funktion des TWM not- wendige Relativbewegung des distal von BAS liegenden Flügelteils ermöglichen. Da der Skle- rit BAS über die HP an den restlichen Flügel mechanisch angekoppelt blieb, war auch die C ek AB --~ ER A | DS, Funktion des TPM gewahrleistet. Die Gelenke c und d durften allerdings nicht zu leichtgangig gewesen sein, da sonst bestimmte Muskeln des TPM (v.a. die indirekten Heber) auf der Hohe des vorderen Gelenkkopfes unwirksam gewor- den waren. Ein Vorteil des kombinierten Systems “TPM a) TPM+TWM” A d) TWM2 ND SLI (I ----MIP1 IT), Cie PE = » ~ Pt «ScH» «Pt4” | Prau: Flugapparat der Libellen 81 + TWM” könnte darin bestanden haben, daß die Flügel (je nachdem, ob der TPM- oder der TWM-Anteil stärker eingesetzt wurde) in un- terschiedlichen Schlagbahnen bewegt werden konnten. Eine stärkere Beteiligung des TWM würde z.B. (infolge der Bewegung des Flügels um die schräg gestellte Achse B) zu einer flache- ren Abschlagsbahn führen. Dabei würde der Flügel gleichzeitig weniger stark proniert; seine Anstellung wäre also der jeweils eingestellten Schlagbahn (bis zu einem gewissen Grad) auto- matisch angepaßt. Jede Weiterentwicklung eines ursprünglichen Antriebssystems “TPM + TWM” muß jedoch zwangsläufig dazu führen, daß die beiden betei- ligten Mechanismen untereinander in Konflikt geraten; sie konnten daher nicht gleichzeitig verstärkt werden, jedenfalls nicht über einen bestimmten Punkt hinaus. Ein vervollkommne- ter TWM, der das Tergum (durch einen kräfu- geren dorsalen Längsmuskel) an der Stelle des hinteren pleuralen Gelenkkopfes weiter nach dorsal wölbt, wäre z.B. auf Kosten des TPM verstärkt: da das Tergum jetzt auf der Höhe des vorderen Gelenkkopfes in der Tierlangsrich- tung beweglicher werden muß (s. z.B. den lin- ken Doppelpfeil in Abb. 24d), müßte seine vor- dere Gelenkverbindung zum Flügel (genauer zum dorsalen Teil des Sklerits BAS) gelöst wer- den; an dieser Stelle würden die Kräfte der TPM-Muskeln schlechter übertragen, die He- belwirkung des TPM wäre also dementspre- chend verkleinert. Umgekehrt kann eine Weiterentwicklung des TPM nur dann stattfin- den, wenn die tergale Kraftübertragung an bei- den Gelenkköpfen, also auch am Gelenk a, ge- steigert wird; eine Vor-Zurück-Beweglichkeit des Tergum müßte somit an dieser Stelle er- schwert werden — der TWM-Anteil würde dem- entsprechend geschwächt. Aus diesen Grün- den tendierte der postulierte Ur-Antrieb “TPM + TWM” wohl schon früh (ab einem bestimmten Punkt, s. unten) zu einer Trennung seiner Teilmechanismen. Er konnte nie über ein Anfangsstadium hinauskommen. Da es nicht wahrscheinlich ist, daß die Muskeln des TPM und TWM in diesem Ur-System gleichzeitig stärker kontrahiert wurden, vermochten die beiden Teilmechanismen sich auch nur bedingt gegenseitig zu unterstützen — bei größerem Krafteinsatz konnten sie wohl höchstens wahl- weise (einzeln) eingesetzt werden (mit dem oben erwähnten Vorteil?). Höhere Schlagge- schwindigkeiten und -frequenzen waren damit aber sicher noch nicht möglich. Nach dem oben Dargestellten bedeutete eine (über das beschriebene Ausgangsstadium hin- ausgehende) Steigerung der Fffektivität des An- triebs also zwangsläufig, daß entweder der TWM- oder der TPM-Anteil reduziert (bzw. modifiziert) werden mußte. Wie konnte es aber überhaupt zu einem System “TPM + TWM” kommen, das sich in gewisser Hinsicht in seiner Weiterentwicklung selbst hemmt? Man kann wohl davon ausgehen, daß die Flügel der frühen Pterygoten primär breit (als vergrößerte “Para- nota”!)) den Thoraxsegmenten ansaßen und zu- nächst, relativ unbeweglich, als Segelflächen fungierten. Bei der sukzessiven Entwicklung der Schlagbeweglichkeit wurde wahrscheinlich anfangs auf jeden dafür in Frage kommenden Muskel “zurückgegriffen”; so wurden Muskeln einbezogen, die ursprünglich für die Beweglich- keit der Segmente vorhanden waren, und auch solche, die der Erzeugung der Atem- und Bein- bewegungen dienten. Das entstandene kombi- !) Der Streit darüber, ob die “Paranota” tergaler, ter- go-pleuraler oder gar rein pleuraler Herkunft sind, soll hier nicht berührt werden. Die Argumente, die Matsuda (1981, S. 387, 388) für eine in einigen Gruppen pleurale Herkunft der Flügel anführt, sprechen z.T. eher für einen tergo-pleuralen Ur- sprung. Abb. 24. Getrennte Ableitung des Flügelantriebs der Odonata (b) TPM), Ephemeroptera (c) TWMI) und Neoptera (d) TWM2) aus einem “Ur-Flugapparat” “TPM+TWM?” (a). Nur der proximale, vordere-mittlere Abschnitt (bis zum Fulcrum-Gelenk) des (aufgeschlagenen) Flügels wurde dargestellt. Pleurum dunkel, Sklerit BAS gekreuzt schraffiert, Membran weit punktiert. Die Sklerit- und Gelenkverbindungen zum Tergum wurden weggelassen (Ausnahmen (c) und (d), wo in der Nähe des Fulcrum liegende Pteralia dargestellt sind — die ver- deckten Sklerite der Flügeldorsalseite Pt1, und Pt1, wurden in (c) gestrichelt angedeutet). Tergum stärker sche- matisiert: für den aufgeschlagenen Flügel ausgezeichnet, für den abgeschlagenen Flügel (außer in (c)) gestrichelt. Der Pfeil am Sklerit BAS deutet die Zugrichtung der “Basalar”-Senker-Muskulatur an, die im Falle von (c) ın Wirklichkeit aus zwei weiter getrennten, am BAS, angreifenden Muskeln besteht. In (b) wurde die Schrägstel- lung der Odonaten-Segmente nicht berücksichtigt; die Membran im Gelenkeinschnitt zwischen BAS und HP ist durchsichtig gedacht, so daß das auf die Dorsalseite gewanderte proximale Gelenk der HP sichtbar wird (vgl. mit a). (c) ist nach den Verhältnissen bei Ephemera gezeichnet; der ursprüngliche Zusammenhang der Sklerite BAS, und BAS, (= BAS) wurde durch punktierte Linien angedeutet. Weiteres s. Text. 82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 nierte System, in dem die fur sich schwach ent- wickelten Teilmechanismen TPM und TWM noch zusammenwirken (sich gegenseitig ver- stärken bzw. ergänzen) konnten, war für den Antrieb der anfänglich kurzen, mit kleiner Am- plitude und niederer Frequenz schlagenden Flügel wohl ausreichend (auch die Verwin- dungsfähigkeit der Flügel konnte auf diesem Stadium der Entwicklung geringfügig sein — die Flügel wurden möglicherweise weitgehend passiv proniert und supiniert). Daß es mit dem “TPM + TWM”-System nicht weiterging, stell- te sich erst später, bei Erreichen seiner Grenzen, heraus. In den folgenden Kapiteln wird der Versuch unternommen, die Grundplankonstruktionen und -funktionen der rezenten Gruppen (*Odo- nata, *Ephemeroptera, *Neoptera) darzustellen und von der postulierten Ausgangskonstruktion “TPM + TWM?” abzuleiten. Dabei soll jeweils auch die Frage verfolgt werden, ob eine “TPM + TWM”-Urform — mit zwei Antriebsmecha- nismen und zwei Schlagachsen — für die Ablei- tung der betreffenden Gruppe überhaupt not- wendig ist, oder ob nicht eher von einer Kon- struktion ausgegangen werden kann, die einem rezenten Flugapparat-Typ ähnlich war. Die Ableitung der Odonata Vergleicht man die in der Abb. 24a dargestell- te hypothetische Ausgangskonstruktion der Pterygoten mit dem Flugapparat der Odonaten (Abb. 24b), so zeigt sich ein wesentlicher Un- terschied: der Flügelteil BAS artikuliert bei Odonaten kaudal an der Stelle e’ (entspricht c4), also innerhalb des Flügels; die Achse B ist da- durch (durch 2 Gelenkpunkte, b und e’) festge- legt (> B’). Diese Situation kann durch eine Verlagerung der Gelenkstelle e nach e’ leicht er- reicht werden — eine nur geringfügig erschei- nende Veränderung des “TPM + TWM”-Sy- stems also, die jedoch eine Verstärkung des TPM und Schwächung des TWM mit sich bringt (der TWM wurde im Zuge der Entwicklung zu einem Stellmechanismus dann weiter um- konstruiert; s. unten). Die daraus folgende Effektivierung des TPM- Anteils ist leicht einzusehen: Durch die Verla- gerung des kaudalen BAS-Gelenks in den Flügel wird der distal von der Achse B’ liegende Flü- gelhauptteil enger an den Skleriten BAS ange- koppelt, und die Wirkung der direkten Senker und indirekten Heber so gleichmäßiger auf die ganze Flugelbreite zwischen den beiden Fulcren verteilt. Bei den rezenten Libellen existieren dementsprechend vordere und hintere direkte Senker, die Basalar- und Subalarmuskeln, die (in Teilfunktionen zwar etwas unterschiedlich) ih- ren Abschlags-Kraftanteil jeweils auf beide Basissklerite des Flügels verteilen; der einzige indirekte Heber, der dvm1, wirkt über ein brei- teres tergales Gelenkgebiet ebenfalls auf beide Teile (möglicherweise sind in der Vorgeschichte der Odonaten mehrere dvm-Muskeln sekundär zusammengerückt). Radioanalplatte und Co- stalplatte (letztere entspricht BAS + HP) der rezenten Odonaten sind damit beim Flügel- schlag als funktionelle Einheit zu betrachten (vgl. auch S. 50). Der Anteil des TWM am Flügelantrieb wird automatisch in dem Moment verringert, in dem der kaudale BAS-Gelenkpunkt e vom Gelenk b weg in den Flügel verlegt wird. Einerseits hängt dies damit zusammen, daß die Funktion des TWM eine nicht-flügelfeste Schlagachse B be- nötigt (vgl. S. 79f.), andererseits wird der dlm (mit der Festlegung der Achse) gleichzeitig zu einem Hilfsmuskel des TPM, da er nun (neben der Bewegung des Flügels um B’) auch den gan- zen Flügel (incl. BAS!) um die TPM-Achse A/B bewegt. Da die tergale Mechanik aber im weite- ren Verlauf der Evolution stärker modifiziert wurde (sie wurde v.a. einer Achse B”=C2/C4 “angepaßt” — die Schubrichtung der dlm konn- te durch Verlagerung von t2 in die Tiefe leicht verändert werden; vgl. auch S. 57ff. und weiter unten), und dabei bestimmte mechanische Vor- aussetzungen für die dlm geschaffen wurden, die deren Wirksamkeit auf einen kurzen Ab- schnitt des Flügelschlags (das Ende der Ab- schlagsphase) einschränken, sind TPM-Antrieb und TWM-Stellmechanismus bei den rezenten Odonaten funktionell (weitgehend!)) getrennt. Die komplizierte tergale Mechanik und die Funktionsbeschränkung des dorsalen Längs- muskels auf einen kurzen Phasenabschnitt “zwischen den beiden Schlagphasen” deuten — bei Vergleich mit den übrigen Pterygoten — dar- auf hin, daß der Vor-Zurückschwingmechanis- mus der Odonaten stark abgeleitet ist. So kann aus der entgegengesetzten Funktion der dorsa- len Längsmuskeln im Meso- und Metathorax bei Zygopteren und Anisozygopteren (vgl. S. 61) auf eine primär in beiden Segmenten gleichartige Funktion geschlossen werden. Ent- gegengesetzt die Flügel bewegende serial-ho- 1) Möglicherweise behielt der dlm bei rezenten Libel- len noch eine geringfügige Abschlags-Teilfunktion (bezüglich der Achse P1/P2 = A/B!) bei. Prau: Flugapparat der Libellen 83 mologe dlm erscheinen ja nur denkbar, wenn die Funktionen zunächst zusätzlich zu einer äl- teren, in beiden Segmenten gleichartigen Funk- tion, nämlich der Abschlagsfunktion, evoluiert wurden; mit der Reduktion der Abschlagsfunk- tion konnten sich die Nebenfunktionen dann zu Hauptfunktionen der dlm entwickeln. Der Vor- Zurückschwingmechanismus kann dabei relativ leicht aus dem ursprünglichen Antriebs-Teilme- chanismus TWM des “Ur-Flugapparates” abge- leitet werden. Es bedarf nur der Versetzung von zwei Gelenkstellen des BAS, um aus der pri- mären Drehachse B des TWM die Achsen B’ und B” (Abb. 24b) zu entwickeln: e muß in den Flügel wandern (> e’), und das distale Gelenk c (zur Humeralplatte) muß auf die Flügel-Dor- salseite verlegt werden (> c2). Durch die Ver- lagerung von c wird zusätzlich zur Achse B’ (P2/C4) eine zweite, in einem stumpfen Winkel zu ihr stehende Scharnierachse B” (C2/C4) ge- bildet. Während B’ (ebenso wie die Achse C1) ungefähr in der Flügelebene liegt und (zusam- men mit C1) die Drehbewegungen des Flügels um die Längsachse im Abschlagsdrehbereich bestimmt, wurde die schräg auf der Flügelfläche stehende Achse B” zur Drehachse des modifi- zierten TWM-Systems (Vor-Zurückschwing- System). Der dlm war möglicherweise in einer Ubergangsphase — bei einem weniger weit “versenkten” Gelenk t2 — auch über die Achse B’ (als Supinator) wirksam. Obwohl wir über die an der Basis der Odonaten abgelaufenen evolutiven Vorgänge wahrscheinlich nie befrie- digende Aufschlüsse bekommen können, er- scheint also prinzipiell folgender Ablauf denk- bar: 1) Der dlm wirkt über B (ursprüngliches TWM-System; Abb. 24a); 2) die Achse B wird zu B’ (der dlm wirkt damit über B’ und A/B); 3) B’ “spaltet” B” ab (der dim wirkt über B’, B” und A/B); 4) t2 wird tiefer gelegt — der dlm wirkt nun (ausschließlich?) über B” (stark mo- difiziertes TWM-System; Abb. 24b). Im Zusammenhang mit der hier im Groben behandelten Evolution des Odonaten-Flugme- chanısmus (Verstärkung des TPM und Speziali- sierung des TWM) stehen zahlreiche (v.a. terga- le) Veränderungen des Skeletts. So wurde das Tergum im Mesothorax auf der Höhe des vor- deren Gelenkkopfes (a) um eine Querachse be- weglich (vgl. S. 58); Tergum und Flügel blei- ben damit an dieser Stelle in engem Kontakt, so daß eine Verschiebung des (vorderen) Tergum in der Tierlängsrichtung, welche den TPM schwächen würde, vermieden wird. Durch ver- schiedene Gelenke im tergalen Seitenbereich (zwischen vTS, hTS und anschließenden Teilen) — die z.T. wohl schon ursprünglich vorhanden waren (vgl. S. 87f.) — wird die Hebelwirkung der dorsalen Längsmuskeln auf das hintere Fulcrum (b) konzentriert; die bei der Kaudal- verschiebung des (mittleren) Tergum stattfin- dende Schrägstellung der dvm wird zu Beginn des Aufschlags (automatisch) wieder rückgän- gig gemacht, so daf der TPM auch hier höch- stens kurzfristig beeinträchtigt wird (vgl. S. 60). Otfensichtlich stehen Antriebssystem und Stellmechanismen bei Odonaten in einem kom- plexen funktionellen Zusammenhang, der im Verlauf der Evolution nur einen schmalen, “gangbaren” Weg zuließ. Dies kann der folgen- de Gedankengang aufzeigen: Setzt man ein Ausgangssystem “TPM + TWM” voraus, so kann angenommen werden, daß die Stelle des Tergalzapfens (TZ) ursprünglich im TWM-Sy- stem (in dem der dlm noch als Senker fungierte) an der Abschlags-Hebelbewegung beteiligt war, und daß das Gelenk t2 noch als längeres (sich in Tierlangsrichtung erstreckendes) Gelenkgebiet zwischen Tergum und Flügel (bzw. Pteralia) ausgebildet war (vgl. auch S. 105). Das Gelenk t2 ist bei rezenten Odonaten jedoch ein punktförmiges Gelenk und ermöglicht erst da- durch die Drehbewegungen im Abschlagsdreh- bereich. Würde das BAS-Gelenk e sich in dieser Situation noch an der Stelle b befinden, so wür- den die Basalarmuskeln pronatorisch wirken (was übrigens primär, bei einem langgestreckten Gelenk t2, nicht der Fall war); da die Gelenk- stelle e jedoch, zur Steigerung der Effizienz des TPM, in den Flügel verlegt wurde (> e’), wurde diese Funktionsänderung vermieden (s. auch S. 50). Daraus ergibt sich, daß der Dreh- mechanismus des Flügels im Abschlagsdrehbe- reich — ebenso wie der Mechanismus des Flügelvor- und -zurückschwingens — erst mit der Weiterentwicklung des TPM überhaupt ent- stehen konnte. Die vorderen Abschlags-An- triebsmuskeln des TPM (bas) machten dabei keinen Funktionswechsel durch (vgl. dazu auch 3 DE Bei den Odonaten führten die Veränderungen im Bereich von t2, der ursprünglich wohl ausge- dehnteren tergalen Hebelstelle des TWM, an- scheinend auch zu stärkeren Abwandlungen der die Hebelbewegung primär übertragenden Ge- lenksklerite (Pteralia 1 und evtl. auch Pterale 4). Die genaue Lage des Pterale 1, das bei Neopte- ren und Ephemeropteren auf der Höhe des hin- teren Fulcrum (bzw. kurz davor) liegt — und 84 sogar sein Vorhandensein bei Libellen über- haupt — ist jedoch umstritten. Keineswegs dürfte z.B. der vordere Tergalsklerit vTS (Abb. la) dem Pterale 1 entsprechen (wie etwa Tan- nert, 1958, annahm), da er sich im Gebiet des vorderen Gelenkkopfes (und der CP) befindet und aufgrund seines Muskels tp mit einem an- deren Sklerit, der Subtegula, homologisiert wer- den kann (vgl. S. 88). Ich vermute stattdessen, daf der vorn an die RAP angrenzende 1. Ge- lenksklerit G1 !) dem Pterale 1 homolog ist. Dar- auf deutet der an diesem Teil inserierende Muskel hca hin, da ein ganz entsprechender Muskel, der ebenfalls in Beziehung zu einem “Pterale 1” steht, bei Neopteren und Epheme- ropteren existiert (vgl. S. 86f.)! Auch die Bezie- hung des Sklerits G1 zur Subcosta (über die Ader cr,; vgl. Abb. 1a) spricht dafür (s. dagegen Matsuda, 1979, l.c. S. 6). Trifft diese Homologi- sierung zu, so hätte das Pterale 1 bei Odonaten seine ursprüngliche Funktion als Übertra- gungselement der Muskelkräfte des TWM voll- ständig verloren. Ein dem Pterale 2 homologer Kutikula-Bereich läge dann innerhalb der RAP, kann jedoch schon deshalb nicht genauer abge- grenzt werden, weil das Pterale 2 erst im Zu- sammenhang mit der Entwicklung der Neopte- rie (Entstehung der Mittelplatten-Gelenke) überhaupt in Erscheinung tritt (vgl. S. 90f.)!). Erschwerend für das Erkennen eines homolo- gen Skleritbereiches ist, daß die RAP bei Libel- len, dort, wo sie dem Fulcrum aufliegt, nicht durchgängig sklerotisiert ist; d.h., Dorsal- und Ventralwand der Flügelbasis sind an der Stelle des Pterale 2 (im Gegensatz zu den Neopteren) voneinander getrennt. Möglicherweise ist dies als eine Folge des abgeleiteten Flügel-Verwin- dungsmechanismus (vgl. S. 47ff.) anzusehen. Erst die Umwandlung des “TPM + TWM”- Antriebssystems in ein weitgehend reines TPM- System ergab (neben der allgemeinen Verstär- kung des Flügelantriebs) eine für die Odona- tenevolution sicher sehr wesentliche Möglich- keit: der mesothorakale Antrieb konnte vom metathorakalen entkoppelt werden, so daß Vor- der- und Hinterflügel verschieden stark oder auch mit einer größeren Phasendifferenz ge- schlagen werden konnten. In einem TWM-Sy- stem existiert dieser, für die Manövrierfähigkeit wesentliche, Vorteil dagegen nicht: Mit vier Flügeln gut fliegende Neopteren (z.B. Locusta) ') Eine Homologisierung des Sklerits G1 mit dem Pterale 2 (Hamilton, 1971) erscheint mir nicht begründbar. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 zeigen, daß ein in beiden Segmenten stärker ent- wickelter TWM-Flügelantrieb nur dann mög- lich ist, wenn das mittlere Phragma das relativ festliegende Ursprungsgebiet beider dlm-Paare bildet; Vorder- und Hinterflügel müssen in die- sem Fall weitgehend synchron bewegt werden, da der Schlagmechanismus der beiden Flügel- paare tergal gekoppelt ist 2). Eine darüber hin- ausgehende Leistungssteigerung des TWM (wie etwa bei Dipteren, Hymenopteren, Strep- sipteren oder Ephemeropteren) konnte anschei- nend stets nur für ein Flügelsegment erreicht werden, was aber auf Kosten des anderen gehen mußte. Der “Verzicht” der Odonaten auf die (bei Effektivierung des TWM in beiden Seg- menten notwendigerweise “segmentkoppeln- den”) dorsalen Längsmuskeln als Antriebsmus- keln kann somit als die wesentliche Vorausset- zung für die (fast vollständige) Unabhängigkeit der Vorder- und Hinterflügel angesehen wer- den. (Erst dadurch wurde andererseits eine in beiden Segmenten unterschiedliche Funktions- differenzierung der dlm möglich.) Der Flügel- antrieb konnte (einmalig bei Pterygoten!) in beiden Segmenten gleichermaßen verstärkt wer- den. Von der Odonatenkonstruktion ausgehend können die Flugapparate der Ephemeropteren (Abb. 24c) und Neopteren (Abb. 24d) — selbst wenn man nur das Antriebssystem berücksich- tigt — nur schwer abgeleitet werden. Zur Her- ausbildung eines TWM-Systems aus dem TPM müßte z.B. der Gelenkpunkt e’ des BAS zu- nächst (unter Schwächung des TPM) zum Ful- crum hin wandern. Eine voll funktionsfähige TWM-Schlagachse B käme aber erst dann zum Einsatz, wenn der Sklerit BAS kaudal direkt bei b artikuliert (dies entspräche der Konstellation 2) Die auch räumlich in engeren Kontakt kommenden Flügel sind bei den Feldheuschrecken allerdings unterschiedlich spezialisiert — die Vorderflügel sind z.B. weitaus weniger an der Auftriebserzeu- gung beteiligt als die Hinterflügel (vgl. Weis-Fogh, 1956, Lc. S. 567). Eine andere Möglichkeit, beide dlm-Paare effektiv einzusetzen, wäre eine gegen- phasische Kontraktion, doch sind mir keine Insek- ten bekannt, die auf dieser Basis eine besser ent- wickelte Flugfähigkeit erreicht haben. Bei weniger gut fliegenden Pterygoten (etwa den Plecopteren, Mecopteren und den meisten Gruppen der Neu- ropteroidea) ist die Kopplung der beiden Thorax- segmente übrigens noch relativ schwach: die bei- den dlm-Paare beeinflussen sich am mittleren Phragma, das einen breiteren ”Gelenk”einschnitt zwischen den Terga darstellt, nur geringfügig. Prau: Flugapparat der Libellen 85 bei Eintagsfliegen). Jetzt konnte (auf dem Weg zu den Neoptera) das Odonaten-Schlagschar- nier a/b (und damit die Schlagachse A/B) redu- ziert werden; daran anschließend könnte der kaudale Artikulationspunkt des Sklerits BAS weiter nach ventral in das Pleurum (> e?) wandern (eine funktionsfähige Schlagachse A/B kann ja vorher nicht einfach “überschritten” werden). Schon für den (zunächst sicher nicht vorteilhaften) “Rückzug” aus dem einen Me- chanismus, zur Erreichung des Ausgangspunk- tes für den anderen, fehlt ein positiver Selek- tionsdruck. Berücksichtigt man die hochspezia- lisierten Stellmechanismen der Odonaten, so erscheint diese Leserichtung noch schwerer vorstellbar. Eine weitere (umgekehrte) Denkmöglichkeit wäre die, daß der Sklerit BAS, ausgehend von einem neopteroiden Zustand (Abb. 24d), erst bei den Vorfahren der Odonaten in den Flügel eingewandert ist. Sie ist aber ebenfalls wenig wahrscheinlich. Eine Drehachse A/B darf z.B. bei dieser Entwicklung nicht zu früh gebildet werden, da sie vom Sklerit BAS sonst nicht mehr “überschritten” werden kann. Ein funk- tionsfähiges Scharniergelenk a/b könnte also erst dann entstehen, wenn die kaudale Gelenk- stelle e” des BAS bereits am Gelenkpunkt b an- gelangt ist (> e). Daraufhin könnte das Ge- lenk e vom Pleurum weg in den Flügel wan- dern; der TWM würde dabei geschwächt, der neugebildete TPM effektiviert. Auch diese Ableitung erfordert also, wenn keine Funk- tionslücke entstehen soll, ein Ubergangsstadium mit zwei Schlagachsen !); sie ist jedoch um- standlicher als der direkt bei “TPM + TWM” beginnende Weg. Ausgehend von einem ein- achsigen System (TWM, Achse B) existiert außerdem kein ersichtlicher Selektionsvorteil, der eine Verlagerung des Sklerits BAS (“zum Zwecke einer spateren Bildung” der Schlagach- se A/B) begründen könnte. Die Ableitung der Ephemeroptera ?) Der Schlagmechanismus der Ephemeropteren (Abb. 24c) läßt sich. ebenfalls leicht vom Ur- Mechanismus “TPM + TWM?” (S. 78ff.) ableiten !) Als unwahrscheinlich kann angesehen werden, daß die Schlagachse B durch Veränderung ihrer Aus- richtung direkt in die Achse A/B übergegangen ist. Außerdem existieren bei rezenten Odonaten an- scheinend noch “Abkémmlinge” der Achse B (B’, B”). Eine ausführliche Darstellung der Flügelmechanik der Ephemeropteren ist in Vorbereitung. N 7 — in diesem Fall durch Weiterentwicklung des TWM und Reduktion des TPM. Die TWM-Ef- fektivitat konnte z.B. durch membranöse Rand- einschnitte auf beiden Seiten des Tergum (Ter- galspalte), wenig kaudal vom Fulcrumgelenk b, gesteigert werden. Da bei den Ephemeropteren aber v.a. der mesothorakale Antrieb weiterent- wickelt wurde — und zwar dadurch, daß der hinter dem Tergaleinschnitt liegende Teil des Tergum (“ScH”, Scutellarhebel) stärker beweg- lich wurde als der davor liegende—, war dies (zwangsläufig, vgl. S. 84) gleichzeitig mit einer weitgehenden Reduktion des metathorakalen Systems verbunden. In dieser Hinsicht besteht eine Analogie zu verschiedenen neopteren Gruppen (vgl. S. 88): Der Hebelsklerit, der die Bewegung des Tergum auf den Flügel überträgt, liegt bei den Ephemeropteren nämlich nicht bei b, auf der Höhe des Fulcrum, sondern kaudal davon (der tergale Hebelpunkt ist also vom pleuralen Drehpunkt weiter entfernt). Er arti- kuliert mit dem Flügel in dem Bereich, in dem bei den Neopteren das Pterale 3 liegt; ein Ptera- le 3 ist zwar bei den Ephemeropteren nicht ein- deutig abgrenzbar, sein Gebiet ist jedoch durch einen zum Fulcrum ziehenden Muskel — den Pterale-3-Muskel der Neopteren, der wohl dem fa der Odonaten homolog ist 3) — gekennzeich- net. Somit ist der Hebelsklerit der Ephemero- ptera möglicherweise dem Pterale 4 der Neo- pteren homolog (und nicht dem Pterale 1) — er soll im weiteren als “Pt 4” bezeichnet wer- den. Bei den Eintagsfliegen führen Bewegungen des Scutellarhebels “ScH” nach oben-vorn- außen und zurück zu einem Ab- bzw. Auf- schlag des Flügels um eine Schlagachse B. Die Achse B wird in diesem Fall v.a. durch ein Scharniergelenk bestimmt, das vor dem “ScH” und “Pt4” und lateral von einem Gelenksklerit Pel, in der dorsalen Kutikula der Flügelbasis liegt (Abb. 24c). Die proximale Kante des Skle- rits Pt1, bildet vorn ein Gelenk zu einem weite- ren kleinen Sklerit Pt1,; das Pt1, artikuliert sei- nerseits, mit seiner medialen Kante, am Tergal- rand (der in der Abb. 24c nicht eingezeichnet ist). Entsprechend der Bewegung des Scutellarhe- bels (mit einer Komponente “nach vorn”; s. Doppelpfeil in Abb. 24c) und der Ausrichtung der Schlagachse B verläuft die Grundschlagbahn des Flügels steil. Sie ist jedoch nicht unverän- derlich. Der Flügel kann nämlich — mitsamt 3) Pm,, bei Brodskyi, 1970. 86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 der Schlagscharnierachse B — durch die am vorderen Teil des Skleriten BAS angreifenden Senkermuskeln (Zugrichtung s. Pfeil links in der Abb. 24c) im Fulcrum-Gelenk b um eine durch b verlaufende Hochachse (s. die senkrecht zu B stehende Punkt-Strichlinie) nach vorn gedreht werden. Je starker diese Muskeln (in Relation zu den dorsalen Langsmuskeln — und zu den Subalarmuskeln, s. unten) kontrahiert werden, um so mehr wird die Achse B wahrend des Abschlags um die Hochachse gedreht, um so horizontaler liegt die resultierende Schwin- gungsebene des Flügels. Dabei werden die Skleri- te Pl, 5, je nach Schlagbahn, mehr oder weni- ger gefaltet, wobei das Pt1, sich mediad über das Pti, bewegt. Ausgespannt begrenzen die Sklerite den Schlagbahn-Spielraum zur Verti- kalen hin. Das ephemeropteroide Antriebssystem kann durch eine Veränderung der Ausrichtung der Schlagachse B (die zu einer vertikaler gestellten Grundschlagbahn führte) sowie durch Ände- rungen der tergalen Mechanik (> “ScH”) leicht von einem “TMP + TWM”-Ausgangs- zustand abgeleitet werden. Auch hier mußten die Stellsysteme wahrscheinlich Hand in Hand mit dem Antriebssystem entwickelt werden: Ei- ne steiler gestellte Grundschlagbahnebene ist z.B. erst dann sinnvoll, wenn der Schlagbahn- Spielraum gleichzeitig zur Horizontalen hin er- weitert wird. Um dies zu erreichen, war es not- wendig, den Flügelteil BAS durch eine Mem- branisierung auf der Höhe des Schlaggelenks a in zwei Elemente zu zerteilen. Vorderer Ge- lenkkopf und Sklerit BAS (bzw. seine beiden entstandenen Teile BAS, und BAS,) wurden auf diese Weise funktionell getrennt; jetzt konnte der (funktionslos gewordene) vordere Gelenk- kopf weiter nach unten abgesenkt werden. Erst mit der Entwicklung des Membranspielraums zwischen BAS, und BAS, und der Reduktion des vorderen Schlaggelenks wurde es überhaupt möglich, die Zugrichtung der vorderen Senker (Basalarmuskeln) stärker zu verändern, d.h. durch eine einfache Verlagerung ihres sternalen Ursprungs schräger zu stellen, so daß sie den Flügel in der oben beschriebenen Weise (mit- samt der Schlagachse B) nach vorn bewegen konnten. Die Basalarmuskeln schlagen den Flügel außerdem wohl weiterhin um die Achse B abwärts. Der Subalarmuskel (nicht abgebil- det) ist anscheinend Antagonist der Basalarmus- keln — zumindest ein Teil des mächtigen Mus- kels (dessen Faserverlauf sehr kompliziert ist) wirkt flügelrückziehend; bezüglich des Flü- gelschlags bleibt er ein synergistischer Senker. Bemerkenswert ist, daß der Flügelteil BAS bei den Ephemeropteren ım hinteren Bereich überhaupt erhalten blieb (als BAS,) und kaudal, wie im ursprünglichen “TPM + TWM”-Sy- stem, zwischen dem Flügel und dem Pleurum schmal endet, ohne dort allerdings ein eigentli- ches Gelenk zu bilden. Anscheinend stellt der Sklerit BAS, ein federndes Element dar, das bei horizontaler geführten Flügelabschlägen senk- recht zur Pleuralebene nach medial abgebogen wird. Da der tergale Hebel “ScH” beim Epheme- ropterenflügel weit kaudal (am “Pt4”) angreift, wurden die davor liegenden Sklerite Pti, und Pel, für die mechanische Führung der Schlag- bahn-Stellbewegung frei. Sie ermöglichen eine Art Faltung des Flügels (s. oben), die den Flügel — im Gegensatz zur Faltbewegung bei Neopte- ren (s. S. 90) — nach vorn führt. Die Schlag-: bahn-Stellbewegung der Ephemeropteren be- trifft außerdem, da die Pteralia 1 ganz proximal liegen, praktisch den ganzen Flügel; bei den Neopteren entstanden die Mittelplatten-Faltge- lenke dagegen weiter distal, innerhalb des Flügels, so daß die proximalen, die Antriebs- kräfte übertragenden Sklerite Pterale 1 und Pte- rale 2 bei einer Schlagbahnänderung nicht mit- bewegt werden (vgl. Abb. 24d). In der Literatur werden die beiden vorderen Pteralia (Pt1, ;) der Ephemeropteren oft mit dem Pterale 1 und 2 der Neopteren homologi- siert (s. z.B. Matsuda, 1970; Hamilton, 1971). Da das Pterale 2 jedoch bei Neopteren dem Fulcrum aufliegt und durch ein laterales Gelenk abgegrenzt ist, das (bezogen auf die Schlagachse B) weiter distal im Flügel liegt als das laterale Gelenk des Pti, der Ephemeropteren (durch welches die Achse B verläuft), ist diese Homo- logisierung wohl nicht zutreffend. Das Pterale 2 entstand — als abgrenzbarer Sklerit — erst in der Stammgruppe der Neoptera, ım Zusammen- hang mit der Ausbildung der Mittelplattenge- lenke und der Neopterie (s. S. 90f.). Das laterale Gelenk des Pterale 1, der Ephemeropte- ra ist daher eher dem lateralen Gelenk des Neopteren-Pterale 1 gleichzusetzen, so daß die beiden Pteralia Pt1, und Pri, also gemeinsam mit dem Pterale 1 der Neoptera homologisiert werden konnen. Fur die Homologisierung (zu- mindest des hinteren Sklerits Pt1,) mit dem Pte- rale 1 der Neoptera spricht übrigens auch ein Muskel (der Pm,, bei Brodskyi, 1970; t-s3 bei Matsuda, 1970), der von einem proximalen Fortsatz des Gelenkstücks nach ventral zu einer Prau: Flugapparat der Libellen 87 Furca-ähnlichen Bildung des Sternalbereichs zieht (Furca bei Matsuda, 1970, l.c. Fig. 43B; Fu bei Brodskyi, 1970, Fig. 4). Da diese “Furca” der Ephemeropteren mit den, nach ventral ver- lagerten, Pleuralarmen der Neoptera homologi- siert werden kann (dafür spricht ein quer-ver- laufender, unpaarer Muskel, der sich — bei Re- duktion der eigentlichen Furca — als Verschmelzungsprodukt der paarigen Furca- Pleuralarm-Muskeln deuten läßt), stimmt der Pt1,-Muskel bezüglich Ansatz und Ursprung mit dem Pterale-1-Muskel der Neoptera über- ein!). (Zur möglichen Homologisierung des Pti-Muskels mit dem hea der Odonaten vgl. S. 83f.) Der Flügelschlagmechanismus der Epheme- roptera (ITWMI1 mit Schlagbahnspielraum “nach vorn”; Abb. 24c) erscheint in seinen Entwick- lungsmöglichkeiten — verglichen mit dem (TWM 2-) Mechanismus der Neoptera — inso- fern “benachteiligt” zu sein, als sich Schlagan- triebs- und Schlagbahnstell-Bewegung zwangs- läufig kaudal am “Pt4” wechselseitig beeinflus- sen. Diese Ausbildung des TWM ist jedoch als so hoch spezialisiert anzusehen, daß sie (einmal entwickelt) als Ausgangspunkt für die Evolu- tion der anderen rezenten Systeme nicht mehr in Frage kommt. Der gegenüber den Neopteren (vgl. S. 88) grundverschiedene Flügelantrieb über das kaudale “Pt4” spricht andererseits auch gegen eine Ableitung des Ephemeropte- ren-Mechanismus aus einem Neopteren-ahnli- chen Vorstadium. Bei einem Vergleich der Ab- bildungen 24c und d könnte man dennoch zu der Ansicht kommen, daß ım Bereich des “Bas- alarsystems” homologe, synapomorphe Ge- meinsamkeiten der beiden Gruppen Neoptera und Ephemeroptera existieren. So könnte die Stelle des (reduzierten) vorderen pleuralen Schlaggelenks der Ephemeropteren ((a)) z.B. mit dem Gelenk f der Neopteren gleichgesetzt werden. Dann wäre der Sklerit BAS, mit dem bas I der Neopteren zu homologisieren, der BAS, mit dem bas II. Dagegen spricht jedoch Verschiedenes: Wie bei den Neopteren existiert auch bei Ephemeropteren — kaudal vom meso- thorakalen Stigma — ein Skleritbezirk zwischen Präscutum und Pleurum, der sog. Tergalarm 1) Bei beiden Gruppen inseriert der Muskel auf der Höhe des Tergalspaltes am Pterale 1,,, (bei Ephe- meropteren dicht beim Vorderrand des Membran- einschnittes). Da die Tergalspalte anscheinend nicht homolog sind (vgl. S. 88), ist diese Uberein- stimmung als Analogie zu interpretieren. (nicht eingezeichnet in der Abb 24; vgl. etwa Weber, 1933; prealare bei Snodgrass, 1935). Diese Struktur steht bei Eintagsfliegen weder mit dem Tergum noch mit dem Pleurum in di- rektem Kontakt und ist nur schwach skleroti- siert (aus funktionellen Gründen: die Basalar- muskeln benötigen Bewegungsspielraum nach vorn!); ventral endet sie unterhalb vom “Rest” des vorderen Gelenkkopfes. Bei den Neoptera liegt die ventrale Kontaktstelle des Tergalarms dagegen dorsal des (neugebildeten) Gelenks f (und spricht so ebenfalls für die auf S. 88f. dar- gelegte pleurale Herkunft des bas 1)2). Die Stel- len f und a und die Sklerite bas I und BAS, wären demnach — wenn man von einer Homo- logie und stabilen Lagebeziehung der Tergalar- me ın den beiden Gruppen ausgeht — nicht ho- molog. Außerdem existiert bei den Ephemero- pteren im Bereich des vorderen Gelenkkopfes noch ein pleuro-tergaler Muskel (t-p10 bei Mat- suda, 1970; Pm,, bei Brodskyi, 1970), wahrend die Muskulatur dieses Gebiets bei den Neopte- ren — im Zusammenhang mit der Bildung des Gelenks f und des Sklerits bas I — anscheinend in das Bewegungssystem der beiden Basalaria übernommen worden ist (vgl. S. 88ff.). Erwähnenswert sind noch einige weitgehende Übereinstimmungen — v.a. im dorsalen Bereich des Sklerits BAS — zwischen Ephemeropteren und Odonaten, die durchaus synapomorph sein konnten (und damit eine nahere Verwandtschaft der beiden Gruppen begründen würden), je- doch hier vorerst als symplesiomorph gewertet werden (d.h.: es wird zunächst angenommen, daß diese Merkmale — in einem noch nicht ge- nauer zu rekonstruierenden Vorzustand — schon bei dem hypothetischen Ur-Flugapparat der Pterygota vorhanden waren). So erstreckt sich vom proximalen Humeralplatten-Gelenk c bei Eintagsfliegen (deutlich etwa bei Ephemera) ein abgrenzbarer kaudaler Teil des BAS in Rich- tung Tergalrand, der mit großer Wahrschein- lichkeit dem Abschnitt phCP der Odonaten- Costalplatte homolog ist (in den Abb. 24a, b, c wurde sein Umriß gestrichelt angedeutet). Pro- ximal davon liegt ein Sklerit, der mit dem (bei Odonaten verschmälerten) Randsklerit RS ho- 2) Es ist anzunehmen, daß der Tergalarm bei Pterygo- ten ursprünglich vorhanden war. Bei Odonaten ist er möglicherweise zu einer Tegula-ähnlichen Struktur umgebildet (s. z.B. die Abb. 4b auf S. 442 bei Tannert, 1958: nicht näher bezeichnete Struk- tur vor der Hinterflügel-CP; und die Ubersichts- abbildung, l.c. S. 439: Struktur frontal-medial von der Vorderflügel-CP). 88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 mologisiert werden kann, da an seinem Kaudal- rand in beiden Gruppen ein homologisierbarer Muskel inseriert (der vca; t-cx4 bei Matsuda, 1970; Pm, bei Brodskyi, 1970)!). Dieser Sklerit steht bei Ephemeropteren seinerseits über ein proximales Punktgelenk mit dem Tergalrand in Kontakt, und zwar kaudal von einem kleinen Sklerit (dem vTS der Odonaten bzw. der Subte- gula der Neopteren), der sich in allen drei re- zenten Gruppen aufgrund seines Muskels (tp, Abb. 2) identifizieren läßt. Durch diese “Auf- hangung” wird der Sklerit BAS, der Eintagsflie- gen (der veränderten Zugrichtung der Basalar- muskeln entsprechend) auch gegenüber dem Tergum nach vorn beweglich und erhält zu- gleich eine Führung und Limitierung. Bei den Neopteren verlief die Entwicklung in diesem Bereich ganz anders, da der Sklerit BAS sich in dieser Gruppe (ebenfalls im Zusammenhang mit einem Funktionswechsel seiner Muskeln) durch Reduktion seiner dorsalen Anteile vollständig vom Tergum gelöst hat; in diesem Fall wurde eine pleurale Aufhängung neu gebildet (durch Abgliederung des bas I vom Pleurum), so daß die BAS- (= bas II-) Bewegung ebenfalls eine “Führung” (in diesem Fall nach medial) erhielt (vgl. unten). Die Ableitung der Neoptera Wie bei den Ephemeroptera ist auch der Flügelschlagmechanismus der Neoptera durch eine Verstarkung des TWM und Reduktion des TPM gekennzeichnet. Im Gegensatz zu den Eintagsfliegen liegen die Membraneinschnitte (Tergalspalte), die zur Effektivierung der He- belbewegung im Tergalrand entstanden, jedoch weiter vorn, etwa auf der Hohe des hinteren pleuralen Flügelgelenkes b. Die tergale Aufwöl- bung und Abflachung wird über das dort lie- gende Pterale 1 auf den Flügel übertragen — auch in den Fällen, in denen (wie bei Hymeno- pteren, Dipteren und Lepidopteren) ein kaudal des Tergalspaltes liegender Scutellarhebel die Hebelfunktion ganz übernimmt. Tergalspalt, Scutellarhebel und Hebelsklerit sind demnach bei Ephemeropteren und Neopteren analoge Bildungen; der TWM beider Gruppen ist unab- 1) Matsuda 1970, Lc. S. 120) gibt für den t-cx4 einer- seits (falschlicherweise) das Pterale 1 (hier Pt1,) als Ansatzpunkt an, andererseits (fir Oligoneuriella und Caenis) einen kleinen, vor dem Pterale 1 lie- genden Skleriten (“supplementary plate”). Viel- leicht wurde der Odonatenmuskel hea (t-cx4') aus diesem Grund als Abkömmling des vca (t-cx4) an- gesehen (l.c. S. 398). hängig von der hypothetischen Ausgangskon- struktion abzuleiten (Abb. 24a>d). (Weitere Argumente gegen eine Evolution des Antriebs- systems der Neoptera aus ephemeropteroiden oder odonatoiden Systemen — oder umgekehrt gegen die Entwicklung der Flugapparate der Eintagsfliegen und Libellen aus Neopteren-ähn- lichen Konstruktionen — sind in den vorherge- henden Kapiteln angeführt.) Die Ableitung des Basalar-Systems der Neopteren erscheint auf den ersten Blick schwieriger als im Falle der Ephemeropteren, da der Sklerit BAS primär, als Bestandteil des Flügels, auf einem Kreisbogen um die Achse A/B “außerhalb” des Körpers (!) bewegt wurde (Abb. 24a und 25a), die Basalarsklerite der re- zenten Neoptera dagegen in den Thorax hinein bewegt werden (Abb. 24d und 25c). Eine konti- nuierliche Transformation der einen Bewe- gungsbahn in die andere (über funktionsfähige Zwischenstadien, d.h. ohne Funktionsstillstand) ist jedoch dann möglich, wenn man annimmt, daß der vordere Gelenkkopf durch die Bildung eines im Pleurum liegenden Gelenkes f in der Vorgeschichte der Neopteren zunehmend nach medial beweglich wurde (Abb. 25a—c). Der vordere Basalarsklerit der Neoptera (bas I) wäre dann als eine pleurale Neubildung, der hintere Sklerit bas II dagegen als Rest des (ursprünglich pteralen) BAS anzusehen (Abb. 24d). Vergli- chen damit erscheinen andere Denkmöglichkei- ten für eine Entstehung der Neopteren-Basala- ria weniger plausibel: Theoretisch könnten z.B. beide Basalarsklerite (bas I und bas II) aus dem Skleriten BAS “herausgeschält” worden sein. Wegen der oben erwähnten Schwierigkeit, die Bewegung des BAS “nach außen” kontinuier- lich in eine Bewegung “nach innen” überzulei- ten, ist dies jedoch wenig wahrscheinlich. Die Muskeln müßten in diesem Fall in komplizierter Weise um die BAS-Kante herum nach innen ge- wandert sein, was ohne drastische Funktions- Veränderung und -Schwächung (Reduktion der BAS-Bewegungsmöglichkeit) nicht vorstellbar ist. (Aus dem gleichen Grund kann auch die Entwicklung des Odonaten-BAS nicht von ei- ner neopteroiden Form mit zwei nach innen be- weglichen BAS(!)-Skleriten ausgegangen sein. Eine Umkehrung der ın Abb. 25a>c darge- stellten Leserichtung — c—a, mit Fortsetzung zum TPM — ist andererseits unwahrscheinlich; s. auch S. 85.) Für die oben dargestellte Neu- bildung des bas I aus dem Pleurum spricht aber auch die komplizierte Muskelausstattung der Neopteren-Basalarıa. Neopteren besitzen näm- Prau: Flugapparat der Libellen 89 “TPM +TWM a b TPM +TWM2 C Abb. 25. Herausbildung des 1. Basalarsklerits (basI) aus dem Pleurum bei Neopteren (Querschnitte durch den vorderen Flügel- und Thoraxbereich). Der gebogene Pfeil gibt die Bewegungsrichtung des proximalen HP-Ge- lenkpunktes c beim Abschlag (Kontraktion des Basalarmuskels) wieder. Pleurum schwarz, Sklerit BAS gekreuzt schraffiert (weitere Sklerite zwischen BAS und Tergum in (a) und (b) weggelassen). HP in (c) nur als Sklerotisie- rung der Flügelunterseite (Verstärkung der Sehne des Basalarmuskels) erhalten. Teg?: Tegula, môglicherweise der letzte (dorsale) “Rest” des BAS. Vgl. auch Abb. 24. lich Muskeln, die von den Basalarskleriten aus an das Tergum ziehen (bei Mickoleit, 1969, die Muskeln 1 und 2; bei Matsuda, 1970, t-p7, t- p9); sie setzen in manchen Gruppen (z.B. Ple- copteren) noch deutlich am vorderen Basalare basl an (ursprünglicher Fall!)!). Weder bei Ephemeropteren noch bei Odonaten sind Mus- keln vorhanden, die von einem zum BAS-Skle- ritbereich gehörigen Teil aus ans Tergum zie- hen; solche Muskeln wären, bei Vorliegen einer Homologie des Flügelabschnittes BAS und der Basalarıa (bas I und II), zumindest für die Ur- form der Pterygota (“TPM + TWM7”) zu for- dern (wo für sie allerdings keine Funktion er- sichtlich ist). Interessanterweise besitzen die Ephemeropteren aber noch einen pleuroterga- len Muskel (t-p10 bei Matsuda, 1970; Pm,, bei Brodskyi, 1970), der unterhalb von (a) — also nicht an einem Teil des BAS-Systems — ent- springt (vgl. S. 87). Wahrscheinlich wurde ein. entsprechender (eventuell homologer) Muskel, der ursprünglich als Verspannmuskel zwischen Pleurum und Tergum vorhanden war, im Ver- lauf des an der Neopterenbasis ablaufenden Prozesses der Angliederung von bas I an bas II 1) Die Beziehung zum basI kann sekundär verwischt sein; die Muskelansätze sind z.B. bei Dipteren auf ein von beiden Basalarskleriten aus ins Körperinne- re ragendes Apodem gewandert. (den Rest des Sklerits BAS) den primar nur sen- kend wirksamen BAS-Muskeln “hinzugefügt” (s. Abb. 25). Im TWM-Antriebssystem der Neopteren verlor der vordere pleurale Gelenkkopf mit der Bildung des Gelenkes f seine Bedeutung — z.T. wohl auch im Zusammenhang damit, daß der vordere Tergalrand in der Tierlangsrichtung ge- genüber den Basalaria beweglicher werden mußte (vgl. S. 81). Dabei kam cs zu einer Re- duktion dorsaler, ursprünglich mit dem Tergal- rand in Beziehung stehender Bestandteile des BAS (und evtl. anderer, vermittelnder Skleri- te?); nur die Tegula blieb anscheinend, als letz- ter Rest, bestehen (Abb. 25c2)). Die auf die Flü- gelunterseite gewanderte (bzw. nur dort skle- rotisierte) Humeralplatte HP überträgt bei Neopteren, wie in den anderen Gruppen, die Zugkräfte der (abschlagenden) Basalarmuskeln auf den Flügel. Sie kann als Sklerit weitgehend 2) Dabei muß hier vorerst offenbleiben, ob die Tegula dem dorsalen Bereich vCP-phCP der Odonaten- Costalplatte entspricht oder dem Randsklerit RS (oder einer bei Odonaten vorhandenen Tegula- ähnlichen Struktur; (vgl. Fußnote S. 872)). Die Sub-tegula (= vTS bei Libellen) gibt sich dagegen in allen drei Pterygotengruppen durch ihre Lage und dem Ansatz des Tergopleuralmuskels (tp) zu erkennen (vgl. S. 87f.). 90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 verschwinden und Sehnencharakter erhalten (z.B. bei Orthopteren). Im ursprünglichen Fall (etwa bei Plecopteren) behielt die Humeralplatte proximal noch eine deutliche Beziehung zum (bei Neopteren vorn sekundar verschmalerten) Sklerit bas II bei (Abb. 24d); in anderen Grup- pen besteht dagegen eine Verbindung zum bas I (wie in Abb. 25c) oder zu beiden Skleriten. Im einzelnen haben die Basalaria (wie auch ihre Muskeln) innerhalb der Neoptera starke Ab- wandlungen erfahren. Reduktion des TPM (und damit des vorderen, pleuralen Schlaggelenks a), Veranderung des BAS-Systems und Verstärkung des TWM wa- ren wahrscheinlich eng miteinander verknüpft. So konnte die Lage des Sklerits BAS (bzw. sei- nes Restes bas II) z.B. erst nach der Reduktion des Schlagscharniers a/b verändert werden; zu einem früheren Zeitpunkt wäre eine Wanderung des kaudalen bas II-Endes zusammen mit dem Gelenk e vom Fulcrumgelenk b nach ventral (—e”) nicht möglich gewesen, da die unter- halb des Sklerits BAS verlaufende Schlagachse A/B nicht überschritten werden konnte. Diese Entwicklungen standen wahrscheinlich weiter- hin in einem Zusammenhang mit der Evolution einer wesentlichen Bewegungsmöglichkeit des Neopterenflügels, der Zurückfaltbarkeit in den Mittelplattengelenken (“Neopterie”). Im Ge- gensatz zu den Ephemeropteren, bei denen der “Scutellarhebel” weiter kaudal angreift, und der Flügel (infolge der Entlastung des Pterale 1 von der Hebelfunktion) nach vorn “faltbar” werden konnte, liegt die tergale Hebelstelle bei Neopte- ren auf der Höhe des Schlaggelenkes b, wo- durch eine ähnliche Entwicklung von vorn- herein verhindert war. Stattdessen wurde in die- sem Fall eine Faltmöglichkeit nach hinten evoluiert. Die dafür wesentlichen Mittelplatten- gelenke (zwischen Pterale 2 und Mittelplatte 1 und zwischen Mittelplatte 1 und Mittelplatte 2; in Abb. 24d gestrichelt) entstanden innerhalb des Flügels, was diese Bewegung, verglichen mit der Faltbewegung der Eintagsfliegen, mecha- nisch unabhängiger vom Schlagantrieb machte. Gleichzeitig wurde die Flügelbasis kaudal-me- dial membranisiert (vgl. Abb. 22) und gab so dem vorn mit der Mittelplatte 1 in Verbindung stehenden Pterale3 den notwendigen Bewe- gungsspielraum. Obwohl über die ursprüngliche Funktion der Neopteren-Basalarıa und ihrer Muskulatur vor- erst (ohne eingehenden Vergleich der verschie- denen Gruppen) noch wenig Sicheres ausgesagt werden kann, ist es wahrscheinlich, daß sie schon früh in einem Zusammenhang mit der Neopterie standen — ohne Veränderung des BAS-Systems (d.h., wenn der Sklerit BAS in der Flügelfläche “liegengeblieben” wäre), wäre die Entwicklung der Neopterie wohl nicht möglich gewesen. Die ın der Abb. 25 rekonstruierte Evolution der Basalarsklerite der Neoptera zeigt, daß die Kraftrichtung der (anfangs reinen) Senkermuskeln des Sklerits BAS (Abb.25a) all- mählich verändert wurde (>b>c: Ent- stehung einer Kraftkomponente nach medial). Die Pterale-3-Muskeln, die den Flügel in den Mittelplattengelenken zu falten vermögen, d.h., nach hinten-innen schwenken, bewegen die (frontal-medial des Falt-Drehpunkts liegende) Gelenkstelle c des Basalarsystems aber zwangs- läufig in die entgegengesetzte Richtung, nach vorn-außen, und dehnen somit die vorderen Senkermuskeln. D.h.: die Muskeln des BAS er- hielten — zusätzlich zu ihrer persistierenden. Funktion als Abschlagsmuskeln — eine Flügel- vorziehfunktion. Die Flügelbasis mußte bei der Entwicklung der Neopterie also nıcht nur ım mittleren bis hinteren Bereich, sondern gleich- zeitig auch vorn stärker verändert werden, da ja primär nicht nur zurück-, sondern auch vor- ziehende Muskeln (wenn dies auch bei einigen rezenten Gruppen nicht mehr zu erkennen ist) beteiligt waren. In der Literatur wird dagegen das Augenmerk hauptsächlich auf die Faltbar- keit des Neopteren-Flügels nach kaudal ge- richtet. Als evolutiver Vorteil dieser Fähigkeit wird immer wieder angeführt, daß sie eine Flügel-Ruhelage über dem Abdomen ermög- licht, wodurch ein geringerer Körperumriß (Tarnung) und eine bessere “Versteckschlüpf- rigkeit” erreicht würden (Feindschutz). Ein Se- lektionsdruck in dieser Richtung kann jedoch anfangs noch gar keine Rolle gespielt haben, da die Fähigkeit zur Flügelfaltung sicher sukzessiv entwickelt wurde und daher zunächst nur ge- ringfügig war. Wahrscheinlich entstand die Neopterie dagegen als eine Möglichkeit, den Schlagbahn-Spielraum zu erweitern (nach vorn, d.h. zu horizontalen Schlagbahnen hin, durch Basalarmuskeln — nach hinten, zu steileren Schlagbahnen hin, durch Pterale-3-Muskeln). Sie diente so primär einer Verbesserung der Manövrierfähigkeit; die Funktion “Ruhefal- tung” (= Extremstellung der Flügel) konnte der ursprünglichen Funktion erst später (als bereits eine weit entwickelte Neopterie vorlag) hinzu- gefügt werden. Da der Flügel bei seiner Faltung nach kaudal gleichzeitig eine pronatorische Drehung Prau: Flugapparat der Libellen 91 durchführt — seine (distale) Vorderkante wird bei der Bewegung in den sich vorn in einem Punkt treffenden Mittelplattengelenken zwangslaufig nach unten bewegt (in der Ruhela- ge ist der Flügel daher maximal proniert) — ist er bei steileren Schlagbahnen automatisch star- ker proniert als bei flacheren. Dies stellte wahr- scheinlich (im Zeitraum der Herausbildung der Neopterie) eine wesentliche, aerodynamisch gunstige Funktionskopplung dar. Bei den re- zenten Neopteren findet sich das Schlagbahn- Stellsystem (= Faltgelenk-, Pterale-3- + Bas- alar-System) jedoch in vielfaltiger Weise abge- wandelt, so daß auch einzelne Funktionen oder Funktionskopplungen sekundär entfielen. Wäh- rend die ursprüngliche Funktion (Veränderung der Schlagbahn) etwa noch bei Hymenopteren, Dipteren und Lepidopteren vorhanden ist, wur- de sie in anderen Gruppen (z.T. nur in einem der beiden Flügelpaare) wieder reduziert (vgl. Pfau, 1977b, 1978a; Pfau & Honomichl, 1979); die Möglichkeit der Ruhefaltung der Flügel über dem Abdomen blieb in den meisten Grup- pen erhalten (Ausnahme Tagfalter). Die als ursprünglich anzusehende phasische Abschlags- Teilfunktion der Basalarmuskeln wurde in meh- reren Gruppen der Neoptera beibehalten, ent- weder zusammen mit der Vorziehfunktion (z.B. Hinterflügel der Caelifera) oder — bei mechani- scher Festlegung einer vorgezogenen Schlaglage des Flügels — als Hauptfunktion (Coleoptera). Sie konnte auch mit anderen (wahrscheinlich neuen) Funktionen kombiniert werden (z.B. im Vorderflügel der Caelifera; vgl. unten). Toni- sche Basalarmuskeln können dagegen als sekun- däre reine Stellmuskeln mit reduzierter An- triebsfunktion, die den Flügel v.a. in einer Schlaglage halten, angesehen werden; Schlag- bahnveränderungen werden in diesem Fall an- scheinend durch die zum Tergum ziehende Basalar-Muskulatur (Muskeln des bas I, s. wei- ter oben) erreicht (vgl. Pfau, 1977 a). Die Basalarmuskeln der Neopteren wurden bisher, aufgrund ihres vor dem Fulcrum liegen- den Zugpunktes, v.a. als Pronatoren gedeutet und als solche für die Pterygoten verallgemei- nert (Snodgrass, 1929, 1935). Eine pronatori- sche Funktion der Muskeln des BAS-Systems war nach dem hier Dargelegten aber ursprüng- lich in keiner der drei Pterygoten-Hauptgrup- pen entwickelt (und damit auch nicht bei der postulierten Ausgangsform “TPM + TWM”). Die Existenz pronatorischer Basalarmuskeln, etwa beim Feldheuschrecken-Vorderflügel (vel. Abb. 22), kann als eine Sonderentwicklung in- nerhalb der Neopteren angesehen werden. Auch bei den Heuschrecken läuft der Flügel- Drehmechanismus jedoch nicht (wie Snodgrass annahm) als Antagonismus pronatorischer Bas- alarmuskeln und supinatorischer Subalarmus- keln, die den Flügel (als Ganzes) um das Ful- crum drehen, ab (einer solchen Bewegung steht bei Neopteren das Tergum/Pterale-1-Längsge- lenk entgegen), sondern als Verwindungsme- chanısmus innerhalb des Flügels. Dafür mußten neue Gelenke evoluiert werden; außerdem mußte die Flügelschlagbahn (durch elastische Mechanismen) festgelegt werden, so daf die ae- rodynamisch ungünstige Kombination einer fla- chen Schlagbahn mit einer (beim Abschlag) ver- stärkten Pronation vermieden wurde (zu Ein- zelheiten der Vorderflügelmechanik der Feldheuschrecken vgl. Pfau, 1977b, 1978a, 1983; Pfau & Nachtigall, 1981). 4. DISKUSSION UND ERGÄNZUNGEN FLÜGELMECHANIK, MUSKELFUNKTIONEN UND AERODYNAMISCHER EFFEKT Einige Muskeln des Flugapparates der Libel- len können als weitgehend reine Muskeln des Schlagantriebssystems, mit nur einer Funktion, angesehen werden. Dies gilt für den 1. Basalar- muskel bas1 (vgl. S. 50) und den 1. Dorsoven- tralmuskel dvm1 (mit einer Einschränkung, vel. S. 60). Auch die zugfederartig wirkenden “Einstellmuskeln” bas2, dvm2 und tp (s.S. 45f.) sind “monofunktionell”. Unter den Muskeln fiir Flügeldrehbewegungen und Schlagbahnande- rungen sind der Fulcroalarmuskel fa (s.S. 54ff.) und der dorsale Längsmuskel dlm (S. 60f.) als weitgehend monofunktionelle Stellmuskeln an- zusehen (der dlm hätte jedoch — wenn er auch außerhalb des hier postulierten Einsatzbereichs kontrahiert wäre — eine weitere Wirkung; vgl. S. 100). Andere Muskeln sind gleichzeitig mehreren mechanischen Systemen zugeordnet (“poly- funktionelle” Muskeln). Bei zwei dieser Mus- keln (subl und sub2; s.S. 50f.) kann man — wenn man die Hebelarme vergleicht — relativ leicht zwischen einer Haupt- und einer Neben- funktion unterscheiden: der 1. Subalarmuskel ist in erster Linie ein Senker (mit supinatori- scher Nebenfunktion), der 2. Subalarmuskel su- piniert den Flügel dagegen hauptsächlich (und ist mit Nebenfunktion Senker). Schwieriger ıst die Unterscheidung von Haupt- und Neben- funktion beim vorderen und hinteren Coxoalar- und beim 3. Subalarmuskel (vca, hca, sub3). Die 92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 beiden Coxoalarmuskeln werden hier (trotz ih- rer deutlichen Aufschlagswirkung) v.a. als spe- zialisierte Supinations- bzw. Pronationsmus- keln der Schlagwendepunkte angesehen (andere Muskeln kommen nicht in Frage bzw. erschei- nen weniger geeignet; vgl. S. 50ff.). Der wohl tonische sub3 könnte als Supinator sowohl beim Aufschlag (S. 56) als auch an der unteren Schlagwendedrehung beteiligt sein (s.S. 98); als Flügelsenker arbeitet er beim Aufschlag außer- dem den Antriebsmuskeln entgegen. Im Libellenflugapparat existieren demnach zwar mechanisch weitgehend unabhängige Be- wegungssysteme für die Schlag- und Stellbewe- gungen (s.S. 34ff.), die vorhandene Muskulatur ist jedoch z.T. zwischen den Systemen angeord- net und bewirkt zwangsweise gekoppelte Bewe- gungen. Im weiteren soll zunächst die Bedeutung der funktionellen Trennung der Be- wegungssysteme allgemeiner diskutiert werden (dabei werden auch die verschiedenen Möglich- keiten zur Beeinflussung der Flügelgeschwin- digkeit erörtert); auf die Funktions-“Über- schneidungen” der Systeme wird dann v.a. in den folgenden Diskussionskapiteln eingegan- gen. An verschiedenen Stellen wird auch die ae- rodynamische Wirkung bestimmter Muskeln diskutiert. Zu diesen (hypothetischen) Folge- rungen muß gesagt werden, daß sie von einer “stationären Aerodynamik” und den dabei übli- chen Kräfteparallelogrammen (vgl. Weis-Fogh & Jensen, 1956; Nachtigall, 1968; Nachtigall in Kaestner, 1972; Dubs, 1979) ausgehen; instatio- näre Effekte, die z.B. bei raschen Anstell- winkeländerungen auftreten können, spielen wahrscheinlich zusätzlich eine nicht geringe Rolle — ihr Anteil kann vorerst höchstens ge- schätzt werden (vgl. z.B. Norberg, 1975). Auch die wechselseitige aerodynamische Beeinflus- sung der Vorder- und Hinterflügel, neuerdings von Azuma et al. (1985) für den langsamen, ste- tigen Steigflug von Sympetrum frequens analy- siert, kann hier keine Berücksichtigung finden. Funktionelle Trennung von Flugmotor und Stellmechanismen Zur Veranschaulichung der Bedeutung von- einander unabhängiger Antriebs- und Stellsy- steme sei zunächst einmal angenommen, ein In- sekt könne nur die Leistung des Schlagantriebs (“Flugmotors”) durch unterschiedlich starke Muskelkontraktionen verändern. Dadurch könnte zwar die Anströmung am Flügel (die sich aus Schlagwind und Fahrtwind ergibt) und auch die am Flügel angreifende Luftkraft (L) verändert werden, jeder Schlaggeschwindigkeit käme aber nur eine bestimmte Luftkraft zu. Vortrieb (V) und Auftrieb (A), in die sich die Luftkraft zerlegen läßt, würden sich damit zwar bei einer Veränderung der Schlaggeschwindig- keit vergrößern oder verkleinern, könnten je- doch nicht unabhängig voneinander variiert werden. Das Tier wäre in seinen Flugfähigkei- ten stark begrenzt, vergleichbar etwa einem ein- fachen Gummimotor-Flugmodell, das bei Kon- stanthaltung des Auftriebs (horizontaler Gera- deausflug) weder beschleunigt noch verlangsamt fliegen kann. Andererseits wäre bei einem Insekt, welches nur die Flügelanstellung aktiv zu variieren vermag, und dessen Schlagan- trieb konstant ist, eine ähnlich eingeschränkte Flugfähigkeit zu erwarten. Sind Flügel-Antrieb und -Anstellung dagegen beide unabhängig voneinander veränderlich — d.h., der Flügel kann in einer bestimmten Anstellung mit ver- | schiedener Geschwindigkeit geschlagen werden und umgekehrt bei einer bestimmten Schlagge- schwindigkeit unterschiedlich angestellt sein — so erweitert sich der Spielraum der Luftkrafter- zeugung beträchtlich. Größe und Richtung der Luftkraft werden weitgehend frei wählbar, die Luftkraftkomponenten A und V dadurch (bis zu einem gewissen Grad) voneinander unabhän- gig. Dieser Gesichtspunkt ist bisher (sicher auch wegen der nur unzureichend untersuchten Me- chanık der Flügel-Stellbewegungen) kaum beachtet worden: Wilson & Weis-Fogh (1962) sahen z.B. bei Schistocerca (Orthoptera) die me- sothorakalen Basalar- und Subalarmuskeln gleichzeitig als Abschlags-“Powermuskeln” und antagonistische Einstellmuskeln der Flügelan- stellung an (gemäß Snodgrass, 1929; vgl. auch S. 91). Sie fanden also bei den direkten Senkern eine Zwangskopplung einer Antriebsfunktion mit einer Stellfunktion vor. Pronation und Supi- nation laufen jedoch im Vorderflügel der Feld- heuschrecken als Flügel-Verwindungen im Flügel, unabhängig von der Antriebsmechanik, ab und können durch einen (weitgehend) schlagneutralen Muskel (den Pterale-3-Muskel = M85) eingestellt werden (vgl. Abb.22 und Pfau, 1977b, 1978a). Auch für die Feldheu- schrecken ergäbe sich demnach (bei Berücksich- tigung der Vorderflügel allein) die Möglichkeit der unabhängigen Variation von Vortrieb und Auftrieb. Vom Antrieb unabhängige Systeme zur Veränderung der Flügelanstellung sind bei den besser fliegenden Pterygoten anscheinend die Regel (s. auch Abb.22; und Pfau, in Vorb.). Prau: Flugapparat der Libellen 93 Libellen haben mehrere Möglichkeiten, die Schlaggeschwindigkeit ihrer Flügel zu steuern: Steigerungen der vom Flugmotor abgegebenen Leistung können z.B. durch kräftigere Kontrak- tion der Antriebsmuskeln und/oder Verstär- kung durch weitere, synergistische Muskeln er- reicht werden. Beim Abschlag steht allerdings nur ein einziger “reiner” Senker zur Verfügung, der basi, während eine Kontraktion des sub1 oder sub2 gleichzeitig eine Flügel-Supination mit sich bringt. Infolge dieser Funktionskopp- lung erscheinen die beiden Muskeln sub1 und sub2 für eine Steigerung der aerodynamischen Abschlagswirkung in doppelter Hinsicht geeig- net — sie erhöhen einerseits die Abschlags- geschwindigkeit (v.a. der subl) und wirken an- dererseits gleichzeitig der (dadurch verstärkten — s.S. 51) passiven Pronation entgegen (v.a. der sub2), wodurch der aerodynamische An- stellwinkel vergrößert oder gleichgehalten wird (vgl. Abb.26a). Theoretisch könnte die Libelle daher entweder mit dem einen direkten (reinen) Senker basi (und mehr oder weniger großem aerodynamischen Anstellwinkel, bestimmt durch den sub2) oder allein mit dem anderen, starken Senker sub1 (der den Anstellwinkel selbst, “automatisch”, groß hält) fliegen. Eine Erhöhung der Geschwindigkeit des aufschla- genden Flügels kann wohl nur durch eine stär- kere Kraftentwicklung im dvm1 erreicht wer- den (der teil-synergistische hca kommt wahr- scheinlich, aufgrund seiner Drehwirkung im Abschlagsdrehbereich, erst am oberen Schlag- wendepunkt ins Spiel — der vca aus entspre- chenden Gründen am unteren; vgl. S. 50f., S. 95 und S. 97f.). Drosselungen des Flugmo- tors sind andererseits durch kleine, tonische Zu- satzmuskeln (“Zugfeder”-Antagonisten) mög- lich (S. 45f.). Eine Verringerung der Flügel- Abschlagsgeschwindigkeit kann durch den dvm2 bewirkt werden; der bas2 (und auch der sub3) wirkt entsprechend auf den aufschlagen- den Flügel. Die Schlagamplitude wird durch diese Muskeln demnach entweder oben oder unten gekürzt; außerdem könnten Phasenver- schiebungen zwischen dem rechten und linken Flügel entweder erzeugt oder ausgeglichen wer- den. Die Wirkung der tp auf die Flügelgeschwindigkeit ist dagegen komplexer. Diese Muskeln beeinflussen sowohl den Aufschlag als auch den Abschlag; sie sind je- weils ın der ersten Schlagphasenhälfte Antago- nist, in der zweiten Synergist der (jeweiligen) Antriebsmuskeln. In welcher Weise dies äußer- lich zum Ausdruck kommt, hängt sicher we- sentlich von der zeitlichen Entwicklung der Kraft in den Powermuskeln ab. Diese Steigerungs- und Drosselungsmöglich- keiten des Libellen-Flugmotors ergeben einen weiten Spielraum der Luftkrafterzeugung nach beiden Seiten hin, einerseits bei der symmetri- schen Krafterzeugung, andererseits auch (bis zu einem gewissen Grad; vgl. S. 43 und S. 45f.) bei rechts-linksseitig asymmetrischen Steuerak- tionen. Das sind Möglichkeiten, die für derartig hoch spezialisierte Lufträuber sicher von größter Bedeutung sind. Veränderung der Flügelanstellung in den beiden Schlagphasen Die beiden mechanisch ganz unterschiedli- chen Drehbereiche des Flügels sind in der mitt- leren Anstellung durch Anschläge voneinander getrennt (s.S. 46ff.). Die Supinatoren des Abschlagsdrehbereichs können daher nicht ın den Aufschlagsdrehbereich “hinüberwirken” und umgekehrt der Pronator des Aufschlags- drehbereichs nicht in den Abschlagsdrehbe- reich. Es ist andererseits (aus energetischen und — da ungünstige Flügelverformungen die Folge wären — wohl auch aerodynamischen Grün- den) nicht wahrscheinlich, daß den Flügel ent- gegengesetzt verwindende Muskeln gleichzeitig aktiv sind, da der Flügel dabei von vorn und hinten unter Spannung gesetzt würde (eine be- reits vorhandene Verwindung würde jedoch nicht rückgängig gemacht). Drehungen in Rich- tung Anstellextrem können demnach erst dann beginnen, wenn die Verwindung im anderen Drehbereich bis zum Anschlag (0°) zurückge- nommen ist. Diese Rückdrehungen im einen Drehbereich, und anschließenden Vorwärtsdre- hungen im anderen, spielen sich an den Schlag- wendepunkten ab und werden z.T. durch spe- zielle Muskeln, z.T. aber auch durch passive Kräfte, beeinflußt (s.S. 97f.). Innerhalb der Schlagphasen werden wahrscheinlich andere Muskeln für Anstelländerungen eingesetzt, wo- bei jedoch für einzelne Muskeln vorerst nicht klar entschieden werden kann, ob sie mehr der Einstellung der Flügelanstellung in der Schlag- phase oder mehr der Beeinflussung der Dreh- geschwindigkeit am Schlagwendepunkt dienen. Für diese Muskeln sollen daher beide Môglich- keiten erörtert werden. Muskeln zur Vergrößerung des aerodynamı- schen Anstellwinkels. — Der 2. Subalarmuskel (sub2) wird wahrscheinlich (in Anbetracht sei- ner gleichzeitigen Abschlagsfunktion) phasisch TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Prau: Flugapparat der Libellen 95 beim Abschlag eingesetzt. Theoretisch könnte er auch phasisch beim Aufschlag kontrahiert werden; er wäre dann aber lediglich ein Syner- gist des bas2 (s.S. 45), da er die Flügelanstel- lung im Aufschlagsdrehbereich wohl nicht zu beeinflussen vermag (s.S. 56)!). Der 2. Subalar- muskel besitzt einen großen Hebelarm zur Drehachse des Abschlagsdrehbereichs und ei- nen kleinen zur Schlagachse. Da bei einer Kon- traktion des Muskels beim Abschlag eine Flü- gel-Supination mit einer Erhöhung der Flügelgeschwindigkeit einhergeht, wird die ae- rodynamische Wirkung des Abschlags in zwei- facher Hinsicht gesteigert (vgl. S. 93 und Abb. 26a). Zusammen mit dem 1. Subalarmuskel ein- gesetzt (bei dem die Abschlagswirkung größer, die Supinationswirkung dagegen kleiner ist als beim sub2), ergibt sich eine Möglichkeit zur Er- weiterung des Bereichs der Luftkrafterzeugung (vgl. S. 92f.). Im Gegensatz zum subi und sub2 erscheint der vca für eine positive Veranderung der Luft- kraft in einem mittleren Abschlagsabschnitt we- niger geeignet zu sein, da er den Flügel gleich- zeitig abbremsen und den Effekt des vergrößer- ten aerodynamischen Anstellwinkels ver- mindern (oder aufheben) würde (vgl. auch S. 51). Der vierte Flügel-Supinator, der sub3, könnte theoretisch (phasisch kontrahiert) den aerodynamischen Anstellwinkel beim Abschlag vergrößern und würde dabei (wie der subl und !) Auf eine phasische Kontraktion des sub2 beim Abschlag deutet auch ein Ausschalt-Experiment von Neville (1960) hin. sub2) gleichzeitig die Schlaggeschwindigkeit erhöhen. Dieser Muskel ist jedoch vergleichs- weise schwach und wird hier — da er mit großer Wahrscheinlichkeit tonisch ist und (als einziger Muskel) den Cubitalsektor supinato- risch zu bewegen vermag — der Aufschlagspha- se und dem Aufschlagsdrehbereich zugeordnet (vgl. S. 56 und weiter unten). Der Fulcroalarmuskel (fa) vergrößert den ae- rodynamischen Anstellwinkel beim Aufschlag. Ein Hinüberwirken in den Abschlagsdrehbe- reich wird als unwahrscheinlich angesehen, da die Muskelwirkung durch den pronatorischen Anschlag des Cubitalsektors begrenzt ist; im Abschlagsdrehbereich wird der RAP-interne Muskel (infolge der beweglichen Aufhängung seines Ursprungs — genau am Gelenk p2 der RAP-Drehachse P2/C4) als Ganzes zusammen mit der RAP bewegt, so daß hier den Muskel dehnende Gegenkräfte fehlen (vgl. S. 55f.). Im Gegensatz zum sub2 des Abschlagsdrehbe- reichs ist der fa schlagneutral. Möglicherweise muß daher bei einer Anderung seiner Kontrak- tionsstärke auch die Schlaggeschwindigkeit des Flügels verändert werden (entsprechend wie beim sub2, der die Geschwindigkeit “automa- tisch” vergrößert); bei Zunahme der fa-Kraft könnte dies z.B. durch dvm1-Verstärkung (oder schwächere Kontraktion des bas2) erreicht wer- den. Muskeln zur Verkleinerung des aerodynami- schen Anstellwinkels. — Der 3.Subalarmuskel (sub3) ist als tonischer Muskel in der Auf- schlagsphase wirksam und beeinflußt die Flügel- Abb. 26. Schemata zur Luftkrafterzeugung. — (a) Vergrößerung des aerodynamischen Anstellwinkels (ß), des Anström-Vektors und der Luftkraftresultierenden (L) bei Kontraktionsverstarkung des sub1 oder sub2 in der Abschlagsphase. Der geometrische Anstellwinkel x wurde hier unverändert belassen. Dies beinhaltet die Annahme, daß (v.a. im Falle einer sub1-Kontraktion) das passiv-pronatorische Drehmoment (hervorgerufen durch den größeren Windfahneneffekt bei größerer Flügelgeschwindigkeit) das supinatorische Drehmoment (bedingt durch die Muskelkontraktion) gerade kompensiert (vgl. auch S. 50f.); (b) Verkleinerung von a und damit verbundene Vergrößerung von ß und L bei Kontraktionsverstärkung des fa in der Aufschlagsphase. Der Anströmvektor wurde unverändert belassen; (c) Verkleinerung von ß, L und Anström-Vektor (bei gleichzeitiger Richtungsänderung desselben — als Folge der Schlagverlangsamung) durch verstärkte sub3-Kon- traktion in der Aufschlagsphase; (d) Möglichkeit der Rücktriebssteigerung bei sehr starker sub3-Kontraktion und Anströmung der Flügelunterseite in der Aufschlagsphase. Dabei wurde angenommen, daß die Luftkraft L, trotz verringerter Anströmgeschwindigkeit, durch die Vergrößerung von ß größer wird. Die Vergrößerung von ß ist sowohl durch die supinatorische Flügeldrehung als auch durch die Richtungsänderung der An- strömung bedingt. Wegen der Erzeugung von Abtrieb ist es fraglich, ob (d) beim Flug zum Einsatz kommt; (e) Zwei Phasen der Vorschwingbewegung (des Anisopteren-Vorderflügels) am Ende des Abschlags. Die Abb. ver- deutlicht die Möglichkeit der Erzeugung von Rücktrieb durch den dlm. x geometrischer Anstellwinkel (Winkel zwischen einer Frontalebene und dem Flügel); ß aerodynamischer Anstellwinkel (Winkel zwischen Luftanströmung und Flügel); L Luftkraftresultierende; S Seitkraft; W Wider- stand. Das schwarze Dreieck an den Flügelquerschnitten kennzeichnet die Vorderkante und Oberseite. 96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 anstellung dort (sobald der 0°-Anschlag des Abschlagsdrehbereichs erreicht ist — vgl. S: 98) durch eine supinatorische Bewegung des Cubitalsektors (s.S. 56). Theoretisch ware der Muskel damit als Antagonist des fa anzusehen; es ist jedoch fraglich, ob er als solcher auch ein- gesetzt wird (d.h., ob er zu gleicher Zeit wie der fa kontrahiert wird), da er den Flugel ja vor allem dann zu supinieren vermag, wenn die pro- natorischen Gegenkrafte gering sind. Der sub3 verwindet den Flügel in Richtung zur aerodyna- mischen Null-Anstellung (tangentiale An- strömung) hin, vielleicht sogar darüber hinaus (> Anströmung der Flügelunterseite). Wäh- rend durch den fa (v.a.) die Vortriebswirkung vergrößert wird (s. Abb. 26b und weiter unten), wird sie durch den sub3 verkleinert — bei gleichzeitiger Verringerung der Aufschlags- geschwindigkeit (vgl. S. 56; Abb. 26c)! Er- reicht der Flügel, bei Anstromung der Flügelun- terseite, negative Anstellwinkel, so kann evtl. sogar Rücktrieb erzeugt werden (Abb. 26d). Da Flügelaufschläge mit wenig oder ohne Vor- triebswirkung (oder mit Rücktrieb) wahr- scheinlich beim langsamen Flug und Rüttel- bis Rückwärtsflug eingesetzt werden, ist zu erwar- ten, daß der sub3 v.a. in diesen Flugsituationen kontrahiert wird (zusammen mit dem dlm? Vgl. S. 100). Bei Libellen liegt damit eine getrennte Ein- stellmöglichkeit der Flügelanstellung in der Ab- und Aufschlagsphase vor, die für die Ma- növrierfähigkeit der Tiere sicher von großer Be- deutung ist. Da die Grundschlagbahnebene des Flügels durch ein Scharniergelenk (p1/p2) fest- gelegt ist, wird der Flügel beim Ab- und Aufschlag in der weitgehend gleichen Bahn be- wegt (vgl. S. 43f.). Daraus ergibt sich für die bei- den Schlagphasen (bei positivem aerodynamı- schem Anstellwinkel) eine ganz unterschiedli- che Ausrichtung der auf dem Flügel stehenden Luftkraftresultierenden (s. Krafteparallelo- gramm der Abb. 26a und b); bei Zerlegung von L in A und V wird ersichtlich, daß beim Ab- schlag mehr Auftrieb als Vortrieb erzeugt wird, beim Aufschlag dagegen mehr Vortrieb als Auf- trieb. Ausgestattet mit der Fähigkeit, die Flügel- anstellung in den beiden Schlagphasen unabhän- gig zu bestimmen, können Libellen daher ent- weder die Auftriebs- oder die Vortriebserzeu- gung betonen (oder natürlich auch beide zu- gleich, und dies entweder rechts-linksseitig sym- metrisch oder asymmetrisch!)) — wiederum ei- ne wesentliche Möglichkeit zur Erweiterung . des Spielraums der Luftkrafterzeugung! Die den aerodynamischen Anstellwinkel ver- größernden Muskeln sub2 und fa besitzen beide (im mittleren Abschnitt ihrer Schlagphase) an- scheinend keinen Muskel-Antagonisten — ıhr “Gegenspieler” ist jeweils die den Flügel (pas- sıv) zur Anströmung hin drehende Luft (vgl. S. 51, 55). Diesem Gesichtspunkt wurde wohl bisher deshalb keine Beachtung geschenkt, weil die Verwindungsmechanik (und -muskulatur) bei keiner Gruppe genauer untersucht war und somit auch Informationen über die genaue Lage der Flügeldrehachse(n) fehlten. Die in der vor- liegenden Arbeit dargestellten Befunde zeigen, daß im Odonatenfligel die Achsen beider Drehbereiche so liegen, daß sich die Haupt- flügelfläche — und damit (wie bei einer Wetter- fahne) auch der aerodynamische Druckpunkt — jeweils hinter der Achse befindet. Aufgrund dieses “Wetterfahneneffekts” wird der aerody- namische Anstellwinkel des Flügels bei Ande- rung der Anströmung gewissermaßen selbst- tätig in einem günstigen Bereich gehalten. Größere Anstellwinkel müssen jeweils aktiv, mit Muskelkraft (gegen die Luft), erzeugt wer- den. In beiden Schlagphasen ist der Flügel ver- wunden, und zwar so, daß er distal stärker pro- niert oder supiniert angestellt ist als proximal. Hierin könnte eine Anpassung an die sich von proximal nach distal ändernde Richtung und Geschwindigkeit der anstromenden Luft gese- hen werden: Da die Flügel-Umfangsgeschwin- digkeit nach distal zunimmt, ändern sich auch Winkel und Stärke der aus Fahrtwind und Schlagwind resultierenden Anströmung zur Flügelspitze hin; dies wird durch die Verwin- dung des Flügels möglicherweise (zumindest zum Teil) kompensiert, so daß der aerodynami- sche Anstellwinkel unterkritisch bleibt. Im Ab- schlagsdrehbereich wird der Flügel deutlich ge- ringer verwunden als im Aufschlagsdrehbe- reich. In Übereinstimmung damit scheint zu 1) Für unilaterale Veränderungen der Flü- gelanstellung scheint v.a. der schlagneutrale fa geeignet zu sein, da bei einer einseitigen Kontrak- tionsänderung der Flügelschlag des anderen Flügels nicht beeinflußt wird. Jedenfalls nicht direkt: wird durch die Korrektur der Flügelanstellung gleich- zeitig die Flügelgeschwindigkeit (passıv, durch die geänderte Anströmung) verändert, so ist natürlich der andere Flügel — indirekt, über die tergale Kopp- lung - mitbetroffen. Prau: Flugapparat der Libellen 97 stehen, daß die Abschlagsgeschwindigkeit des Flügels kleiner ist als die Aufschlagsgeschwin- digkeit (vgl. den Film von v. Holst, 1950; Nachtigall in Kaestner, 1972; Savage et al., 1979, nach Daten von Norberg, 1975). Da das AusmaR der Flügelverwindung von zahlreichen Kräften abhängt (von den Stellmus- keln und von der Luftanströmung direkt — von den Antriebsmuskeln, welche über die Verände- rung der Flügel- und Fluggeschwindigkeit die Anströmung verändern, indirekt), läßt das äußerliche Bild eines mehr oder weniger ver- wundenen Flügels (Abb. 27) für sich keine Rückschlüsse auf den Kontraktionszustand der Muskeln zu. Es ist außerdem noch völlig offen, in welcher Weise sich der Flügel genau bei un- terschiedlichen aktiven und passiven Kräften verwindet (und sonst verformt). Hier können in Zukunft nur experimentelle Untersuchungen, die zahlreiche Faktoren berücksichtigen, eine weitere Aufklärung erbringen. Schlagwendepunkte Norberg (1975) kam zu dem Ergebnis, daß ein großer Teil der Auftriebskräfte beim Libel- len-Rüttelflug von den Wendepunktsdrehungen des Flügels herrührt. Der aerodynamische Me- chanismus ist jedoch bisher weitgehend unbe- kannt. Möglicherweise wird der sog. “flip me- chanism” (Weis-Fogh, 1973) genutzt!). Savage et al. (1979) wiesen dagegen auf andere Mecha- nismen hin: die Autoren zeigten anhand von Modellexperimenten, daß bei rüttelfliegenden Libellen die Supination der unteren Schlagwen- depunktsdrehung — über Wirbelbildung bzw. Sogwirkung an der hinteren und vorderen Flügelkante — einen großen Anteil an der Auf- triebserzeugung hat. Aktive Wendepunktsdrehungen haben wohl zumindest die Bedeutung, daß der Flügel mög- lichst rasch in eine der Abschlags- oder Auf- schlagsanströmung “angepaßte” Anstellung ge- schwungen wird und so schon von Beginn an in der Schlagphase Luftkräfte erzeugen kann — ein rein passives Umschwingen würde wahr- scheinlich Verluste mit sich bringen. Anderer- seits kann die Flügeldrehgeschwindigkeit durch eine spezialisierte “Wendepunktsmuskulatur” variiert werden, so daß Phasenunterschiede der Flügel beider Seiten erzeugt (oder ausgeglichen) werden können; dies dürfte für Steueraktionen von Bedeutung sein. Für die Pronation der oberen Schlagwende steht den Libellen (nach Ausschluß des bast — vgl. S. 50) nur der im Abschlagsdrehbereich pronatorische, wahrscheinlich phasisch-aktive !) Dabei könnten elastische Verspannungen im Flügel (z.B. bei früher Kontraktion des hca) eine Rolle spielen. Ein in beiden Hälften der Schlagwende- drehung den Flügel entgegengesetzt verwindender, einen bistabilen Effekt erzeugender Muskel (vgl. Pfau & Nachtigall, 1981: Subalarmuskel im Vor- derflügel von Locusta) war bei Libellen jedoch nicht nachweisbar. Abb. 27. Männchen von Aeshna cyanea im Rittelflug. Man beachte den stark supinatorisch verwundenen, aufschlagenden rechten Hinterflügel sowie den weit vorgeschwungenen, am Abschlagsende befindlichen rech- ten Vorderflügel. 98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 hea zur Verfügung (s.S. 44, 50). Der Muskel besitzt zusatzlich eine Aufschlagswirkung. Am Auf-Abschlags-Umkehrpunkt kontrahiert, be- wirkt er demnach — da er auch die Aufschlags- bewegung des Flügels fortsetzt und die obere Amplitude vergrößert — für sich wohl keine wesentlich beschleunigte Pronationsdrehung!). Schnellere Drehbewegungen des Flügels sind deswegen aber nicht ausgeschlossen; sie könn- ten durch einen (gegenüber der hca-Kontrak- tion) verfrühten Einsatz der Abschlags-An- triebsmuskeln erreicht werden. Dabei würde der aufschlagende Flügel zunächst abgebremst und dann abgeschlagen; der sich (spät) kontra- hierende hca würde weniger schnell verkürzt (isometrische Kontraktion), so daß der Flügel rascher (“auf der Stelle”) proniert würde. Wahr- scheinlich wird der Muskel sub2 erst nach dem hea, innerhalb der Abschlagsphase, kontrahiert (vgl. S. 93ff.); bei fruhzeitigem Einsatz könnte er allerdings ebenfalls an der Wendepunktsdre- hung beteiligt sein — er würde in diesem Fall den Drehwinkel und die Drehgeschwindigkeit (im Abschlagsdrehbereich) verkleinern. Der vca wird hier als Supinator der unteren Schlagwende betrachtet (s.S. 51). Da der Mus- kel (der wie der hca medial von der Schlagachse am Flügel angreift) zusätzlich eine Aufschlags- funktion besitzt, bremst er den Flügel bei seiner Kontraktion ab (im Gegensatz zum hca, s. oben). Der Flügel wird dadurch schnell, prak- tisch auf der Stelle, bis zum 0°-Anschlag des Abschlagsdrehbereichs hin supiniert (die Mus- kelwirkung wird wohl dadurch noch gesteigert, daß der vca bis zum Ende der Schlagphase durch die Abschlagsbewegung gedehnt wird — dies steht wiederum im Gegensatz zum hca). Eine verlängerte Wendedrehung (bei gleichzei- tig weıter fortgesetztem Flügelschlag — wie sie der hca allein bewirken kann, s. oben) kann durch einen schwächeren Einsatz des vca und wohl auch durch stärkere (bzw. späte) Kontrak- tion von Abschlagsmuskeln erreicht werden. Der Muskel sub3 vermag (als tonischer Muskel) die Supination des unteren Schlagumkehrpunk- tes fortzusetzen (Zugfederwirkung mit begin- nendem Aufschlag; vgl. S. 56). In diesem Fall wären die Kräfte zweier gleichsinnig drehender Muskeln aus verschiedenen Drehbereichen (vca, 1) Im Insekten-Flugfilm von v. Holst (1950) zeigt der linke Vorderflügel der Libelle (anscheinend Aeshna juncea L.) eine solche, einen größeren Abschnitt der Aufschlagsphase einnehmende “verlängerte” pronatorische Wendepunktsdrehung. sub3) hintereinandergeschaltet. Würde der Su- pinator des Abschlagsdrehbereichs (vca) da- gegen nicht kontrahiert, so würde der sub3 — infolge seines Hebelarms zur Drehachse P2/C4 — zu Beginn des Aufschlags auch den ersten Teil der Drehung übernehmen. Für den Muskel fa muß ebenfalls eine Beeinflussung der unteren Wendepunktsdrehung erwogen werden: der Muskel würde (bei tonischer Kontraktion oder im Falle eines frühen phasischen Einsatzes) ent- sprechend wie der sub2 (vgl. oben) den Dreh- winkel und die Drehgeschwindigkeit des Flügels (in diesem Fall im Aufschlagsdrehbe- reich) verkleinern. Setzt der Muskel fa dagegen phasisch erst sehr spät ein, so wäre er an der oberen Wendepunktsdrehung — wiederum nur in dem zum Aufschlagsdrehbereich gehörigen Teil der Drehung — pronatorisch beteiligt. Demnach besitzen die Odonaten mit den Muskeln hca und vca eine spezialisierte Musku- latur für die Flügeldrehbewegungen des oberen und unteren Schlagwendepunkts. Da außer die- sen Muskeln aber noch weitere beteiligt sein können (fa, sub2, sub3), und auch passive Kräf- te sicher eine nicht unwesentliche Rolle spielen, kann im einzelnen mit sehr komplexen Kraftbe- ziehungen und vielfältigen Bewegungsablaufen gerechnet werden. Funktionsmorphologische und experimentel- le Untersuchungen weisen neuerdings auch für andere Insektengruppen darauf hin, daß den Schlagwendepunktsdrehungen (oder zumindest einer von beiden, also entweder der pronatori- schen oder der supinatorischen Drehung) eine größere Bedeutung zukommt. Bei Locusta ist z.B. die Pronation des oberen Schlagumkehr- punktes in vielfältiger und komplexer Weise ak- tiv beeinflußbar (Pfau, 1977b, 1978a, 1983; Pfau & Nachtigall, 1981, s. Fußnote S. 97). Hier spielen drei Muskeltypen (Basalarmuskeln, Sub- alarmuskel und Pterale-3-Muskel) eine Rolle und können, je nach Kontraktionskraft und -zeitpunkt, die Drehbewegung modifizieren; die als wesentliche “Initiator”-Pronatoren ein- gesetzten Basalarmuskeln sind in diesem Fall gleichzeitig starke Senker des Flügelantriebssy- stems (vgl. dazu auch S. 91 und Abb. 22). Die Supination des unteren Schlagwendepunkts läuft dagegen bei Locusta (und auch bei Cetonia und Geotrupes; vgl. Pfau & Honomichl, 1979) in relativ einfacher Weise, weitgehend passıv, ab: sowie der Flügel langsamer wird und pro- nierende Kräfte abnehmen, schwingt er ın eine elastisch bedingte supinierte Grundanstellung zurück. Demgegenüber sind die Dipteren (Cal- Prau: Flugapparat der Libellen 99 dim — 7. DI ip Nam j Ne wet a) b) C) Abb. 28. Grundschlagbahnebene und Möglichkeiten zur Schlagbahn-Veränderung beim Vorderflügel der Aniso- pteren (a,b) und der Zygopteren + Anisozygopteren (c). In (a) und (b) wurde die postulierte “Vorbereitung” der Vorschwingbewegung — durch Kontraktion des vca vor dem dlm (s. S. 60f.) — illustriert; (a) Schwache Kontraktion des dlm (evtl. zusätzlich schwache vca-Kontraktion) > großer Schlagbahnwinkel X, am Ende des Abschlags (der Flügel schwingt etwa in der Fortsetzung der Grundschlagbahn aus); (b) Starke Kontraktion des dim (und vca) > kleiner Winkel À, am Abschlagsende; (c) Schema der Wirkung des dim und pa bei Zygopteren und Anisozygopteren (vgl. S. 61). liphora) anscheinend wie die Libellen in der La- ge, auch am unteren Umkehrpunkt des Schlags einen Wendepunktsmuskel (Supinator) einzu- setzen (pt4, in Abb. 22); bei rein tonischer Kon- traktion wurde dieser Muskel allerdings — ent- sprechend wie der sub3 der Odonaten — erst zu Beginn des Aufschlags wirksam (vgl. Pfau, in Vorb.). Die bisherigen Befunde zeigen also, daß bei den Pterygoten mehrere, ganz unterschiedli- che Wendepunkts-Mechanismen evoluiert wur- den. Veränderung der Schlagbahn Das Flügel-Vorschwingen ist von anderen Bewegungen abhängig (vgl. S. 57ff.). Es kann z.B. erst dann beginnen, wenn der Flügel sich in einer bestimmten Anstellung (0° = “zwischen” den Drehbereichen) befindet. Ist er proniert an- gestellt, muß dem Vorschwingen demnach eine Supination vorausgehen; ein Uberlappen der Vorgänge würde beide Bewegungen “schwer- gängig” (weniger effektiv) machen. Die Funk- tion der tergalen Hebelkette ist andererseits von der sich im Schlagablauf verändernden Ausrich- tung der Scharnierachse C2/C4 abhängig und kann erst in der unteren Hälfte des Schlags “ablaufen”; vorher ist ein vollständiges Vor- schwingen des Flügels, bis hin zum Anschlag der dhCP an der phCP, nicht möglich. Beim Aufschlag schließlich verhindert der starke indi- rekte Heber dvm1 ein Vorschwingen — ein (phasischer) Einsatz des dim kann daher in die- ser Phase als unwahrscheinlich angesehen wer- den. Der Kontraktions-Zeitpunkt der dorsalen Längsmuskeln läßt sich also auf den unteren Abschlagsabschnitt, anschließend an eine supi- natorische Drehung bis 0° (vca), eingrenzen; da- bei ist zu erwarten, daß die durch den dim be- wirkte Schlagbahnanderung je nach der Kon- traktion des Supinators verschieden ausfällt (vgl. S. 60f. und Abb. 28a, b). Dieses (aus der Mechanik erschlossene) zeitlich enge Zusam- menwirken der beiden Muskeln dlm und vca er- klärt möglicherweise einen Irrtum Neville’s (1960): Neville beobachtete, daß die Flügelba- sisplatten RAP und CP am Abschlagsende aus- einanderweichen und schrieb dies (aufgrund der etwa gleichzeitig stattfindenden, die vca-Kon- traktion anzeigenden Bewegung des Rand- sklerits RS nach ventral) einer Kontraktion des Muskels vca zu (wobei er sich allerdings z.T. widersprüchlich äußert — s. Anm. 17, S. 116). 100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 Abb. 29. Flügel-Vorschwingen bei Aeshna cyanea (nach einer Sequenz des Films von G. Rüppell, 1981; Tier in Großaufnahme, Abdomenende außerhalb des Bildes). Eineinhalb aufeinanderfolgende Schläge des linken Vor- derflügels eines rüttelfliegenden Männchens sind dargestellt: W Abschlag 1, A Aufschlag 1, @ Abschlag 2 (Flügelspitzenbahn). Da das Tier danach eine Roll- und Gierbewegung durchführte, konnte der folgende Aufschlag nicht mehr verfolgt werden. Der Flügelumriß wurde nur für beide Abschlags-Endpunkte (für densel- ben Vorderflügel) ausgezeichnet, der Hinterflügel wurde weggelassen. Die Pfeile kennzeichnen den jeweiligen Beginn der Supinationsdrehung. Die Abstände zwischen den letzten vier Punkten des 2. Abschlags sind per- spektivisch verkürzt, da der vorschwingende Flügel schließlich nach innen-oben (vom Betrachter weg) aus- schwingt. Die in diesem Fall einseitig (links) durchgeführte Steueraktion wirkte anscheinend einem Wegdrehen des Tiers nach rechts entgegen (die dorsalen Langsmuskeln dim sind bei nur ein-seitig weit abschlagendem Flügel v.a. auf der Seite der großen Amplitude wirksam; vgl. S. 58). Schlagfrequenz ungefähr 31 Hz; Bildfre- quenz 460 Bilder/sec. Der dlm, der ein Auseinanderweichen der Basisplatten allein zu bewirken vermag, wurde von Neville jedoch nicht berücksichtigt. Die aerodynamische Wirkung einer am Abschlagsende zunehmend horizontal und dann auch nach oben führenden Flügelbahn könnte ın einer Veränderung der Ausrichtung der Luft- kraftresultierenden L, die mehr und mehr nach oben und dann auch nach hinten zeigt, gesehen werden (Abb. 26e). Dadurch würde zunehmend Rücktrieb erzeugt, was bedeuten würde, daß der dim beim Abbremsen des schnellen Vor- wärtsflugs oder beim Rüttel- und Rückwärts- flug eingesetzt wird; die in der Abb. 30 nach Schnappschüssen umgezeichneten Momentauf- nahmen freifliegender Tiere, sowie Abb. 29, ei- ne Szene aus einem Libellenflugfilm von Rüp- pell, können als Belege dafür angesehen werden. Das Flügel-Vorschwingen ist, aufgrund der schräg zur Flügelfläche stehenden Achse C2/C4, mit einer supinatorischen Bewegungs- Komponente verknüpft, die dafür sorgt, daß die Anstellung im Verlauf der Vorschwing-Bewe- gung allmählich verändert wird (vgl. Abb. 26e und 28 und S. 59). Dies könnte als eine auto- matische Anpassung an die sich gleichzeitig än- dernde Richtung der Luftanströmung interpre- tiert werden. Das auf das Vorschwingen folgen- de Zurückschwingen des Flügels soll hier nicht näher untersucht werden; in diesem Fall ist die weniger stabile Flügelhinterkante “führend”. Für die Beurteilung des daran anschließenden (eigentlichen) Aufschlags ist wesentlich, ob der dim phasisch kontrahiert wurde oder tonisch ist. Ein tonischer dim würde beim Aufschlag (v.a. zu Beginn, da die dlm-Wirkung später “gesperrt” ist) dem dvml antagonistisch gegen- uberstehen (vgl. S. 60; Abb. 9e, f); er wurde die Autschlagsgeschwindigkeit verringern, da ein Teil der dvm1-Kraft gegen den dlm aufge- wandt werden müßte. Auf einen Schlagab- schnitt mit Rucktriebserzeugung (Vorschwin- Prau: Flugapparat der Libellen 101 Abb. 30. Eierlegende Sympetrum striolatum — langsamer Manövrierflug auf engem Raum (nach geblitzten Aufnahmen). Beim oberen Paar zeigt das Männchen, beim unteren das Weibchen weit vorgeschwungene Vor- derflügel. gen) würde also eine Phase mit verminderter Vortriebswirkung (Aufschlag) folgen. Verglichen mit Zygopteren und Anisozygo- pteren (vgl. S. 61f.) sind die Anisopteren sicher darın als abgeleitet zu betrachten, dag Verande- rungen der Schlagbahn nur beim Vorderfligel stattfinden können; nur der Mesothorax besitzt einen kräftigen dim, nur beim Vorderflügel ist im Gelenk c2/c4 ein weites Vorschwingen mög- lich. Im Metathorax ist der dlm bis auf ein win- ziges Längsmuskel-Rudiment, das an einem sehr kurzen Hebelapodem angreift, reduziert; das Metatergum ist außerdem kaum verform- bar, die Beweglichkeit des Flügels im c2/c4-Ge- lenk stark eingeschränkt. Der Muskel pa fehlt bei Anisopteren in beiden Segmenten. Bei den Zygopteren und bei der Gattung Epiophlebia können dagegen Vorder- und Hinterflügel eine Vor-Zurückschwingbewegung ausführen, der Hinterflügel allerdings mit einer stark abwei- chenden Mechanik und einem gegenüber dem Mesothorax “umgekehrten” Muskelantagonis- mus. In beiden Thoraxsegmenten stehen pro Flügel (den dvm1 nicht gerechnet) zwei antago- nistische Muskeln zur Verfügung (dlm und pa). Dies deutet darauf hin, daß bei diesen Libellen der Schlagbahnwinkel A der Vorder- und Hin- terflügel — ausgehend von einer gegenüber An- isopteren mehr horizontal stehenden Grund- Schlagbahnebene (vgl. S. 43f.; Abb. 28) — nach beiden Seiten hin verändert werden kann. Eine Vergrößerung des Winkels würde mehr Vor- trieb erbringen, eine Verkleinerung mehr Auf- trieb bis Rücktrieb. Beim Anisopteren-Vor- derflügel, dessen Grund-Schlagbahnebene stei- ler steht, ist dagegen nur eine Verkleinerung von À möglich (vgl. auch S. 109f.)!). SENSORISCHE KONTROLLE UND FLUGSTEUERUNG Über die Sinnesorgane des Flug-Steuerungs- systems der Libellen existieren bisher nur weni- ge experimentelle Untersuchungen. Sie betref- fen v.a. Rezeptoren aus nicht-thorakalen Kör- perbereichen, z.B. Rezeptoren im Halsbereich und Augen (Mittelstaedt, 1950), Windrezepto- ren am Kopf (Sveshnikov, 1973) und Antennen (Gewecke et al., 1974). Im folgenden sollen einige Uberlegungen zur funktionellen Bedeutung der hier untersuchten Flügel-Mechanorezeptoren angeschlossen wer- den. Da diese bis jetzt nur auf die funktions- anatomische Analyse und relativ wenige elek- trophysiologische Ableitungen (v.a. Summen- ableitungen) gegründet werden können, müssen sie als vorläufig angesehen werden. Für weitere Aufschlüsse sind v.a. Untersuchungen an ein- !) Der Muskel fa kann nicht als ein Antagonist des dim betrachtet werden, da er beim Vorschwingen (ähnlich wie beim Flügelschlag oder bei der Prona- tion im Abschlagsdrehbereich, vgl. 55f.) als Gan- zes zusammen mit der RAP bewegt wird (auch die Auslenkung des Fulcrum — vgl. S. 58 f. — dehnt den Muskel höchstens minimal). Der fa ist bei Anisopteren außerdem im Metathorax (in dem der Vorschwing-Mechanismus reduziert ist!) normal entwickelt. 102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 zelnen Sensillen notwendig; auch Ableitungen an vor dem Windkanal fliegenden Tieren müßten durchgeführt werden. Die im Kapitel 2 (S. 62ff.) dargestellten Be- funde zur Beanspruchung der Mechanorezepto- ren lassen darauf schließen, daß sowohl das Chordotonalorgan als auch die beiden Reihen campaniformer Sensillen für die Kontrolle der Flügeldrehbewegungen um die Längsachse ein- gesetzt werden; andere Bewegungen zeigen dagegen keinen Einfluß auf die Rezeptoren, je- denfalls keinen direkten. Beide Rezeptorsyste- me werden in beiden Drehbereichen zugbean- sprucht, das CH bei den Drehungen von den Anstellextremen (S,,,, oder pmax) zur mittleren Anstellung (0°) hin, die Kutikula der CF1,2-Ge- biete dagegen bei Flügeldrehungen in die entge- gengesetzte Richtung (CF1: 0° nach p,,,,; CF2: 0° nach S,,,,). Die elektrophysiologischen Un- tersuchungen ergaben, daß im Falle der campa- niformen Sensillen anscheinend nur diese Zug- beanspruchungen der Kutikula (in Langsrich- tung der Kutikulargruben) zur Erregung führen, während für das CH sowohl Dehnun- gen als auch Entdehnungen reizwirksam sind. Das deutet (ebenso wie auch das stark unter- schiedliche Erregungsmuster der CH- und CF- Sensillen) darauf hin, daß sich die beiden Rezep- torsysteme funktionell ergänzen (jedoch nicht auf einfache Weise, d.h. nach Drehbereich und Drehrichtung). Die Ableitungen weisen auf eine unterschied- liche Spezialisierung der etwas 50 Einzelscolo- pidien des Chordotonalorgans hin. Da das Or- gan in den beiden Drehbereichen nicht symme- trisch beansprucht wird (s. Abb. 16, 17), könnten die verschieden großen Dehn-Ent- dehngeschwindigkeiten zur Erregung unter- schiedlicher Einzelsensillen führen und damit eine sensorische Trennung der Drehbereiche ermöglichen. Vielleicht sind die Drehbereiche aber auch dadurch getrennt, daß der ventrale CH-Ansatz im Abschlagsdrehbereich ın einer etwas anderen Ebene bewegt wird als im Aufschlagsdrehbereich (vgl. Abb. 17a), so daß — je nach Ausrichtung — nur ganz bestimmte Einzelscolopidien oder Scolopidialbündel ge- dehnt bzw. entdehnt werden!). Weiterhin kann man annehmen, daß durch die Dehnungen an- dere Sensillen erregt werden als durch die Ent- dehnungen (u.U. eine zweite Sinneszelle im sel- ben Scolopidium?!)), wodurch auch der Dreh- sinn kodiert wäre. Einige Ableitungen deuten sogar darauf hin, daß bestimmte Sensillen (oder Sensillengruppen) nur in relativ kleinen Winkel- abschnitten innerhalb der Drehbereiche anspre- chen (Abb. 19f) oder nur durch ganz bestimmte Dehn- bzw. Entdehngeschwindigkeiten erregt werden (s. die “on-off”-Spikes in den Abb. 18c—e und 19b). Zur Aufklarung der unterschiedlichen Sensil- len-Spezialisierungen müßten Untersuchungen durchgeführt werden, die auch die komplizierte Transformation des Reizes zum CH berück- sichtigen, d.h., die tatsächliche jeweilige Einwir- kung auf das CH in Rechnung stellen. Selbst bei äußerlich einfachen Reizen kann aus der Dreh- geschwindigkeit ja keinesfalls direkt auf die Dehn- bzw. Entdehngeschwindigkeit des Re- zeptors geschlossen werden (s. Abb. 17b). So wird die Flügel-Drehbewegung im Abschlags- drehbereich nur stark untersetzt auf den (klei- neren!) Hebel vCH weitergegeben — sie wird nicht 1:1 in die für die CH-Beanspruchung we- . sentliche CoS-Bewegung umgesetzt, da Flügel und RAP auch als Ganzes bewegt werden (vgl. S. 47ff.). Elastische Kräfte wirken zusätzlich modifizierend; die Flügelverwindungen werden z.B. zu den Extremen p,,,, und S,,,, hin zuneh- mend durch in der Kutikula entstehende Ge- genkräfte erschwert (s.S. 49f., 53f.). Aus die- sen Gründen ist zu erwarten, daß das CH (un- ter der Voraussetzung einer konstanten Drehkraft) bis p,,,, relativ langsam und gering- fügig entdehnt wird und von da an, bis 0° zurück, eine etwas schnellere Dehnung (bis hin zum Längen-Maximum) erfährt. Die Entdeh- nung im Aufschlagsdrehbereich ist wohl an- fangs noch schneller, wird aber dann zum Ver- windungsextrem So, hin wieder langsamer; hier ist das Organ am Endpunkt stärker ver- kürzt als bei pax im Abschlagsdrehbereich. Bei der Zurückdrehung zur 0°-Anstellung beschleu- nigen dann wieder, wie im Abschlagsdrehbe- reich, elastische Krafte. Nicht nur die Drehbe- reiche waren demnach verschieden (“asymme- trisch”), sondern in ihnen jeweils auch die Dehn- und Entdehnvorgänge der Hin- und Zurückdrehung. An den Stellen maximaler äußerer Geschwindigkeit der Sinusdrehungen (0°, Abb. 17b) gehen die Längenänderungen des 1) Eine elektronenmikroskopische Untersuchung (Risler in Vorb.) ergab inzwischen, daß die Scolo- pidien in verschiedenen Bündeln zusammengefaßt vorliegen. Es zeigten sich dabei auch morphologi- sche Unterschiede zwischen den Scolopidien, die meist nur eine, in einigen Fällen aber auch zwei Sinneszellen enthalten. Prau: Flugapparat der Libellen 103 Rezeptors jeweils auf Null zurück. Für die natürlichen 0°-Durchgänge der Schlagwende- punkte ist außerdem zu erwarten, daß dort auch die Drehkräfte klein werden; die Anschläge zwischen den Drehbereichen bedingen ja, daß verschiedene Muskeln nacheinander kontrahiert werden müssen, wenn die ganze Drehung aktiv vollzogen werden soll!) (vgl. S. 46f.). Môgli- cherweise sind dies Eigenschaften des Systems, die zur Schonung des bei 0° maximal gedehnten CH beitragen. Die stark phasischen Sensillen des Chordoto- nalorgans sind (nach dem oben Dargestellten) nicht nur in der Lage, anzuzeigen, dal Anstell- winkeländerungen stattfinden, sondern könnten darüber hinaus auch Informationen über Dreh- bereich und -richtung sowie Geschwindigkeit und Dauer liefern. Damit würden sowohl Anderungen der Kontraktion der Drehmuskeln (Pronatoren, Supinatoren) als auch Änderungen der Luftanströmung (auftretende Wirbel beim Abreißen der Strömung etc.!?) registriert. Diese Informationen des CH sind im gedehnten Zu- stand des Organs (um 0°) anscheinend beson- ders genau — hier werden die meisten Impulse abgegeben, das Organ reagiert auch äußerst empfindlich. Möglicherweise ist der 0°-Durch- gang der Schlagwendepunkte demnach ein we- sentlicher Bezugspunkt für die Muskeleinsätze. Da die schnellen Schlagwendepunktsdrehungen des Flügels andererseits wie es scheint an der Luftkrafterzeugung beteiligt sind (Norberg, 1975; Savage et al., 1979), wäre ihre sensorische Kontrolle auch direkt für die Steuerung der Luftkräfte von Bedeutung. Genauere Messun- gen der Anstellung im 0°-Bereich könnten außerdem beim Segeln wesentlich sein (s. weiter unten). Die große Bedeutung der Flügel- Chordotonalorgane für den Flug konnte durch Ausschaltexperimente, die an frisch gefangenen Tieren ım Freiland durchgeführt wurden, de- monstriert werden: Bei diesen Versuchen wur- den die CH bei verschiedenen Großlibellen (Aeshna cyanea, Anax imperator, Orthetrum cancellatum) in allen vier Flügeln durch Abtren- nung von ihren ventralen kutikulären Ansatz- Stiftchen außer Funktion gesetzt. Die mit einer feinen Nadel vorgenommenen Eingriffe, die nur winzige äußere Verletzungen (fast ohne Austritt von Gewebsflüssigkeit) mit sich bringen, hatten deutliche Veränderungen des Flugvermögens zur Folge: entweder stürzten die Tiere beim ') Dieser Gesichtspunkt wurde in der Abb. 17b nicht berücksichtigt. Abflug sofort in Spiralen zu Boden oder sie wa- ren (zwei von insgesamt sechs Exemplaren) nur noch zu einem langsamen, stetigen Steigflug in der Lage. Die langen Reihen der dicht aneinander- schließenden campaniformen Sensillen auf der Oberseite der RAP, über die bei den Flügel- drehungen von 0° nach Smax bzw. Pmax Zugspan- nungen anscheinend von proximal nach distal hinweglaufen (s.S. 64ff. und S. 72ff.), könnten genauere Informationen über Drehbereich, Drehablauf und geometrische Flügelanstellung (im ganzen Winkelbereich) liefern. Wahrschein- lich werden die einzelnen Sensillen nacheinan- der und jeweils nur in einem kleinen Winkel- abschnitt der Drehung erregt. Der Drehbereich würde durch die betroffene Reihe (CF1 oder CF2) angezeigt, die Geschwindigkeit der Dre- hung durch die Abfolge des Erregungsbeginns der Sensillen, wobei das zuletzt gereizte Sensil- lum den erreichten Winkelort wiedergeben würde). Bei den Rückdrehungen zur 0°-An- stellung werden die (nur bei Drehung nach p,,,,, bzw. Smax erregten) Sensillen wohl in umgekehr- ter Folge “abgeschaltet” — mit ähnlichen Mög- lichkeiten des Informationsgewinns. Das (im Gegensatz zum CH) phasisch-tonische Muster der Einzelsensillen enthält jedoch wahrschein- lich noch weitere Informationen. So bleiben die den Drehvorgängen und erreichten Anstellun- gen entsprechenden Sensillenerregungen für ei- ne bestimmte Zeit “stehen”, wodurch statische Anstellungen — Flügelanstellungen beim Segeln (s. weiter unten) oder auch die Ruhe- und Start- bereitschafts-Anstellung (s.S. 56) — perzipiert werden könnten. Darüber hinaus ist evtl. sogar die Messung der aktiven und passiven Kräfte möglich, die — je nachdem, ob sie von proximal oder distal aus, oder auch von beiden Seiten (entgegengesetzt oder gleichsinnig), wirken — unterschiedliche Zugspannungen in der RAP erzeugen’). Ausgangs- und Endzustand und zeitliche Entwicklung der bei einer Flügel- drehung auftretenden kutikulären Spannungen (und der sie verursachenden Kräfte) würden 2) Das (schmale) CH erscheint dagegen für genaue Winkelmessungen der Flügelanstellung wenig geeignet. 3) Dadurch könnten z.B. auch Überbeanspruchungen der Kutikula angezeigt werden: bei stärkerem Wind stellen Libellen, v.a. die grofsflügligen Aesh- nıden, das Fliegen ein oder weichen in ruhigere Zo- nen aus. 104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 sich im Verhalten der einzelnen Sensillen und im Gesamtverhalten der Reihen ausdriicken (vgl. dazu auch Pfau, 1983, S. 75). Die große Anzahl und enge Stellung der Sensillen und ihre wahrscheinlich überlappenden Bereiche deuten dabei auf einen großen Dynamikbereich für ge- ringe Anstellwinkel- bzw. Kraftänderungen und auf eine hohe Meßgenauigkeit hin. Proxi- mal sind beide Sensillenreihen deutlich verbrei- tert (Abb. 11, 12). In dieser Region, die hier dem 0°-Anstellbereich zugeordnet wird, liegen die meisten Sensillen; sie adaptieren anschei- nend weniger rasch als die distal liegenden Sen- sillen der extremen Anstellungen (vgl. S. 73). Dies könnte für eine genauere (feiner gestufte) Registrierung mittlerer Flügelanstellungen (und ihrer Kräfte), etwa beim Segelflug, sprechen; die längere “Gebrauchszeit” der Information wür- de damit übereinstimmen. (Die extremen Flügelanstellungen, p oder S,,,., treten da- gegen sicher nur innerhalb der Schlagphasen auf und sind — verglichen mit statischen Anstellun- gen des Segelflugs — von kurzer Dauer.) Außerdem sind — ähnlich wie beim CH — im 0°-Bereich der Schlagwendepunktsdrehungen besonders genaue Messungen möglich. Funktionsmorphologische und elektrophy- siologische Untersuchungen an Locusta haben ergeben, daß auch hier die Flügeldrehbewegun- gen durch Dehnungsrezeptoren (im Mesotho- rax ein Streckrezeptor und ein Chordotonalor- gan) perzipiert werden können (vgl. Pfau, 1978b, 1983). Beide Rezeptoren sind in einer mittleren Flügelanstellung maximal gedehnt (für den Streckrezeptor konnte dies durch Längen- messungen und elektrophysiologische Ablei- tungen belegt werden). Die Dehnungsrezepto- ren des Vorderflügels von Locusta stimmen also in dieser Hinsicht mit dem CH der Libellen auf- fallend überein (der Streckrezeptor feuert allerdings nur bei Dehnung) — beide sind aber mit Sicherheit nicht dem Libellen-CH homo- log! Für die Felder campaniformer Sensillen in der Flügelbasis anderer Pterygoten ergab sich aus funktionsmorphologischen Untersuchun- gen, daß sie ebenfalls als Meßsysteme der Flügelanstellung eingesetzt werden könnten (vgl. Pfau & Honomichl, 1979: verschiedene Felder in der Flügelbasis von Cetonia und Geo- trupes; Pfau, 1983: Sensillen in der ventralen Basis der Subcosta von Locusta!)). Ein elektro- physiologischer Nachweis war in diesen Fallen jedoch noch nicht méglich. EVOLUTION Evolution der Flugapparate und Cladogenese der Pterygoten Die Flügel-Antriebssysteme der drei Ptery- goten-Hauptgruppen konnten zwanglos (d.h. über funktionsfähige Zwischenstadien) von ei- ner Ausgangskonstruktion abgeleitet werden, welche die beiden rezent verwirklichten An- triebsprinzipien TWM und TPM noch in sich vereinte (Kap. 3; Abb. 24). Von diesem Urflug- apparat “IPM+TWM” ausgehende Effektivie- rungen des Flügelantriebs erscheinen nur mög- lich, wenn entweder der TWM- oder der TPM- Anteil seine Antriebsfunktion verliert, das ver- bleibende System dagegen weiterentwickelt wird. Damit einhergehend konnten einzelne Be- standteile des von der Antriebsfunktion “befrei- ten” Teilsystems — in allen drei Pterygotenli- nien — fur die Entwicklung von Stellmechanis- men genutzt werden. Sie wurden wohl (da eine Antriebssteigerung nur dann vorteilhaft ist, wenn auch die Manövrierfähigkeit verbessert wird) gleichzeitig und in wechselseitiger Ab- stimmung mit dem jeweiligen Antriebssystem evoluiert. Bei der Reduktion (bzw. Transforma- tion) des einen Teilmechanismus, und der Ef- fektivierung des anderen, wurde in allen drei Entwicklungslinien nur eine der beiden ursprünglichen Schlagachsen in das neue An- triebssystem einbezogen, die andere wurde ent- weder aufgegeben oder (modifiziert) in das Stellsystem übernommen. Odonata. — Bei den Odonaten wurde der TPM-Flügelantrieb weiterentwickelt und der TWM-Anteil reduziert; Vorder- und Hinter- flügel konnten damit unabhängig werden. Der Vor- und Zurückschwingmechanismus ging (mit unterschiedlichem Ergebnis in beiden Seg- menten) aus dem TWM hervor; er ist damit — ebenso wie der Flügelverwindungsmechanismus im Abschlagsdrehbereich (s.S. 83) — als eine Autapomorphie der Odonaten zu betrachten. Dagegen handelt es sich beim Verwindungsme- chanismus des Aufschlagsdrehbereichs môgli- cherweise um eine Plesiomorphie. 1) Gettrup (1966) konnte bei Schistocerca einen Ein- fluß entsprechender (in der Subcosta-Basis liegen- der) Sensillen des Hinterflügels auf die Anstellung des Vorderflügels nachweisen (intersegmentaler Reflex). Prau: Flugapparat der Libellen 105 Ephemeroptera. — Die Ephemeropteren ef- fektivierten den TWM, bei gleichzeitiger Re- duktion des TPM. Da der TWM hier weit kau- dal am Flügel angreift und zu einer steilen Grundschlagbahnebene führt, mußte in dieser Gruppe gleichzeitig eine Flügelbeweglichkeit nach vorn (für flachere Schlagbahnen) entwik- kelt werden: Die Zugrichtung der Basalarmus- keln wurde verändert; ım Zusammenhang damit wurde die (ursprünglich einheitliche) Unterseite des Sklerits BAS zweigeteilt — dabei verloren beide Teile (BAS, und BAS,) ihre Verbindung zum vorderen Gelenkkopf. Die vor dem ver- bliebenen Schlaggelenk b (und proximal vom tergalen Schlagscharnier) liegenden Pteralia 1 konnten als Führungselemente der Schlagbahn- Stellbewegung eingesetzt werden. Neoptera. — Die Neopteren beschritten ebenfalls den Weg der TWM-Weiterentwick- lung; die tergale Hebelstelle des Flügels wurde jedoch weiter vorn (auf der Höhe des Fulcrum) ausgebildet. Da ein effektiver Flügelschlag bei dieser Anordnung eine geringere Hebelbewe- gung des Tergum als bei den Ephemeropteren erfordert, war die Entwicklung einer mehr hori- zontal stehenden Grundschlagbahn der Flügel von vornherein begünstigt. Im Zusammenhang damit (und wohl auch mit der Reduktion des TPM, welcher primär steilere Schlagbahnen ermöglichte; vgl. S. 80f.) wurde die Neopterie, die Beweglichkeit des Flügels nach kaudal, zur Erzeugung steilerer Schlagbahnen evoluiert. Das BAS-System des “TPM+TWM” wurde (durch Einbeziehung eines pleuralen Elements und Reduktion der tergalen Verbindungen) stark abgewandelt; die Antriebsmuskeln des BAS (die vorderen direkten Senker) erhielten ei- ne zusätzliche Funktion als Vorziehmuskeln im Schlagbahn-Stellsystem. Für die drei rezenten Pterygotengruppen er- geben sich drei verschiedene systematische Gliederungsmöglichkeiten (vgl. etwa Hennig, 1969). Es wurde der Versuch unternommen, synapomorphe Gemeinsamkeiten der Flugappa- rate zu finden, die eine nähere Verwandtschaft zweier Gruppen belegen könnten. Die Schlag- und Stellsysteme erwiesen sich jedoch in den drei Gruppen als grundsätzlich verschieden und alternativ — es war nicht möglich, zwei Grup- pen von einer nur ihnen gemeinsamen Aus- gangsform abzuleiten oder Gründe dafür zu fin- den, daß ein rezenter Flugapparat- -Typ selbst als präadaptiv für die Entwicklung eines anderen anzusehen ist. So spricht z.B. gegen die zu- nächst naheliegende Ableitung der Ephemero- ptera und Neoptera (beide mit TWM-Antrieb) von einer nur ihnen gemeinsamen Ausgangs- form, daß in beiden Gruppen unterschiedliche Gebiete des Tergalrandes für die Flügelhebe- lung spezialisiert sind!). Es ist daher anzuneh- men, daß der TWM-Anteil ursprünglich (beim Ausgangsmechanismus “TPM+TWM”) noch relativ schwach entwickelt und uneffektiv war; die Hebelzone des Tergum für den TWM war wahrscheinlich zu Beginn noch langgestreckt, so daß beide Entwicklungslinien (und auch die zu den Odonaten führende Linie) unabhängig davon ausgehen konnten. Dagegen war die Rekonstruktion eines allen drei Gruppen gemeinsamen “Urflugapparates” durchführbar. Daraus kann man den Schluß zie- hen, daß die effektiveren Flugapparate (TPM, TWM1 und TWM 2; Abb. 24) in den drei Pte- rygotenlinien getrennt (unabhängig) entwickelt wurden; d.h., die Evolution ging in den einzel- nen Stammeruppen jeweils von einem noch ähnlichen System (“TPM+TWM?”) aus. Das stark umstrittene Problem der stammesge- schichtlichen Aufspaltung der Pterygoten (vgl. etwa Hennig, 1969; Kristensen, 1975; Matsuda, 1981) erscheint demnach mit Hilfe des Flugap- parates nicht lösbar. Die Möglichkeit, alle re- zenten Flugapparate von einer Ausgangskon- struktion abzuleiten, spricht aber andererseits für die Monophylie der *Pterygota (*vgl. Fußnote S. 76f.), und somit dafür, daß die Flug- fähigkeit der Insekten nur einmal entstanden ist! Die von Matsuda (1981) vorgebrachten Argu- mente für eine polyphyletische Entstehung der Pterygota und ihrer Flugfähigkeit beruhen großenteils auf Voraussetzungen, die (nach den hier vorgelegten Ergebnissen) einer Korrektur bedürfen (vgl. dazu auch S. 35f.). In seiner ablehnenden Einstellung gegenüber Verfechtern monophyletischer Gruppenbildungen und hy- pothetischen Rekonstruktionen von Ahnformen übersieht Matsuda auch, daß selbst di- oder po- lyphyletische Systeme, falls sie sich begründen lassen, in jeder einzelnen Linie wieder mono- phyletisch sind, was bedeutet, daß jetzt Rekon- 1) Die Konvergenz der beiden TWM-Systeme läßt sich mit Hilfe der Hebel-Pteralia (“Pt4” bzw. Pr) begründen: die Sklerite liegen verschiedenen Berei- chen der Flügelbasis an und sind durch direkt oder in ihrer Nähe ansetzende, nicht-homologe (!), sn gekennzeichnet (s. auch S. 85 und S. 86f.). 106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 struktionen von mehreren “hypothetical com- mon ancestors” notwendig werden. Die Argu- mente Matsuda’s (auf die ich im weiteren einge- hen möchte) vermögen eine polyphyletische Entstehung der Pterygota jedoch nicht zu sı- chern, so daß die Frage nach den verwandt- schaftlichen Beziehungen der Pterygoten- Hauptgruppen (also nach der Abfolge der er- sten Aufspaltungen) immer noch offen ist. Im Gegensatz zur Ansicht Matsuda’s existie- ren keine prinzipiellen Schwierigkeiten, die Flugapparate der Odonaten, Ephemeropteren und Neopteren von einer gemeinsamen Aus- gangskonstruktion abzuleiten. Die Merkmale, die Matsuda zur Begründung einer getrennten Evolution der Odonata aus Machiliden-ahnli- chen Vorfahren anführt (1981, S. 391), stellen, wie er selbst weiß (lc. S. 391 unten), vorerst rei- ne Ähnlichkeiten dar; es konnten keine Argu- mente dafür angeführt werden, daß sie echte Synapomorphien sind, also auf einen nur diesen beiden Gruppen (den Machiliden und Odona- ten) gemeinsamen Vorfahren zurückgehen. Das als besonders wesentlich erachtete Merkmal “large compound eyes” (l.c. S. 391) erscheint z.B. wenig überzeugend (nicht überzeugender jedenfalls als etwa die borstenförmige Fühler- geißel der Palaeoptera, s. Hennig, 1969); andere Merkmale (“four intratergal apophyses”, “pseu- doprescutum”, “median lobe of labium”) müs- sen — ım Hinblick auf ihre Homologie und Verbreitung — überprüft werden (vgl. dazu auch Kristensen, 1975). Die von Matsuda aut Seite 390f. aufgeführten, die Odonaten allein auszeichnenden Merkmale des Flugapparates (die nach seiner Ansicht für eine unabhängige Entwicklung der Libellen sprechen) sind nur zum Teil als autapomorph anzusehen, so etwa das Merkmal “Synthorax”!) oder auch das Merkmal Nr. 2: “dorsal extension of the mes- episternum...”. Dagegen beruht die Annahme, daß nur bei den Odonaten zwei “Humeralplat- ten” existieren, auf nicht korrekten Homologi- sierungen (s. dazu auch S. 41), bzw. darauf, daß offensichtlich nicht damit gerechnet wurde, daß ein Teil des Basalare ursprünglich im Flügel “inkorporiert” vorgelegen haben könnte. Über das Vorhandensein eines dem Pterale 1 homolo- gen Sklerits äußert sich Matsuda widersprüch- 1) Zum sog. “Synthorax” der Odonata muß ange- merkt werden, daß die Verschmelzung der beiden Flugsegmente nur die Pleura und Sterna betrifft — die Schlag- und Stellsysteme der Flügelpaare sind funktionell unabhängig! lich (vgl. 1981, S. 391 oben und S. 392 oben; oder 1970, 1979)2). Das Fehlen des 2. und 3. Axillarsklerits bei Odonaten erscheint anderer- seits nicht verwunderlich; diese Sklerite sind erst mit dem Entstehen der Neopterie in “typi- scher” Ausprägung zu erwarten — sie fehlen damit bei Odonaten und Ephemeropteren von vornherein (vgl. S. 83f., 86f., 90). Die von Mat- suda weiter angeführten Autapomorphien sind entweder problematisch (die hintere Kappen- sehne und das Fehlen des Subalare; vgl. dazu S. 41) oder können, wie die doppelte pleurale Flügelartikulation (vgl. S. 78ff. und 82ff.) oder das Vorhandensein bzw. Fehlen verschiedener Muskeln, als Plesiomorphien angesehen wer- ‘den. Zur Beurteilung der Thorax-Muskulatur der Odonaten äußert sich Matsuda allerdings nur vage. Einige dieser Muskeln sind (bei Berücksichtigung der in der vorliegenden Ar- beit begründeten neuen Homologievorstellun- : gen) nicht mehr als Odonaten-autapomorph an- zusehen, bei anderen ist die Homologie noch unklar (dies betrifft auch einige der auf $. 393 bei Matsuda aufgeführten, die Ephemeropteren und Odonaten unterscheidenden Muskeln und Muskelfunktionen, die jetzt durchaus anders in- terpretiert werden können). Der Verlust mehre- rer Muskeln bei Odonaten erscheint mir übri- gens nicht verwunderlich — er wird im Zusam- menhang mit der Effektivierung und Okonomisierung des TPM-Flugapparates ver- ständlich (auch innerhalb der Neoptera kam es in mehreren Linien zu Vereinfachungen und Reduktionen!); im einzelnen muß aber noch un- tersucht werden, welche Muskeln tatsächlich bei den Odonaten fehlen. Für eine gemeinsame Evolution der Epheme- ropteren und Neopteren (in einer unabhängigen Linie) sprechen nach Matsuda mehrere Merk- male, die alle als im Zusammenhang mit der Entwicklung der Flügel “neu” entstandene Strukturen gedeutet werden. Auch diese Merk- male sind jedoch — in einem anderen Licht be- sehen — ganz anders interpretierbar: Die vor- genommene Homologisierung dreier Axillar- sklerite bei Ephemeropteren und Neopteren (Punkt (1), lc. S. 392) ist keineswegs gut begründet; die Axillarsklerite 2 und 3 fehlten bei den Ephemeropteren wohl schon primär (s. oben). — Die als Punkt (2) aufgeführte Pleural- leiste (mıt dem Fulcrum und dem darauf ruhen- den “2. Axillare”) ergibt sich als eine Symple- 2) Zur möglichen Homologie des Sklerits G1 und des Pterale 1 vgl. S. 83 f. und S. 86 f. Prau: Flugapparat der Libellen 107 siomorphie der Pterygoten. — Die Epipleurite Basalare und Subalare (Punkt (3)) sind wohl kei- ne Synapomorphien der Ephemeropteren und : Neopteren, sondern ebenfalls (partiell) symple- siomorph; das BAS-System der Ephemeropte- ren (dessen vorderer Abschnitt bisher als “Basalare” bezeichnet wurde) kann dabei nur mit einem Teil der Basalaria der Neopteren, dem 2. Basalare (basII), homologisiert werden (vgl. S. 85f. und S. 88f.). — Die nach Matsuda als synapomorph zu wertende Differenzierung der “notal wing processes” (Punkt (4)) erscheint mir schwer belegbar. Da der Flügelantrieb “über einen Scutellarhebel” bei Neopteren (Hy- menopteren, Dipteren) als eine späte Errungen- schaft (innerhalb der Neoptera) anzusehen ist, kann zumindest dieser spezielle Antrieb als eine Konvergenz der Neoptera und Ephemeroptera betrachtet werden; er läuft bei den Ephemero- pteren über das “Pt4” (= posterior notal pro- cess; Matsuda, 1970), bei Hymenopteren und Dipteren dagegen über das Pti und den “ante- rior notal process” ab. — Im Punkt (5) wird die Thoraxmuskulatur der Ephemeroptera und Neoptera als weitgehend synapomorph gewer- tet. Auch diese Übereinstimmungen sind wohl großenteils Symplesiomorphien (einzelne Mus- keln sind wahrscheinlich auch noch bei den Odonaten vorhanden). — Der mit starken dor- salen Längsmuskeln versehene Flügelantrieb (Punkt (6)) unterscheidet sich schließlich bei den Ephemeropteren und Neopteren grund- sätzlich (zur Konvergenz der TWM-Systeme bei Ephemeropteren und Neopteren vgl. S. 85 und 88). Eine der Grundlagen der Argumentation Matsuda’s ist noch kritisch zu betrachten. Der Autor schließt aus dem Alter der “Odonata” (die dabei wohl im weiteren Sinne verstanden werden, also nicht als *Odonata) und anderer a Gruppen (“older than Palaeodictyopte- a”) auf eine konvergente Entwicklung aus en Vorfahren (l.c. S. 390). Fossil re- präsentierte “Taxa” sind jedoch prinzipiell nur schwer als monophyletische Einheiten zu si- chern; ihre Zuordnung zu den rezenten Grup- pen ist meistens problematisch und umstritten (vgl. dazu die grundlegenden Gedanken von Hennig, 1969, 1982, oder Schlee, 1971). Die Folgerung Matsuda’s erscheint mir daher nicht sehr gewichtig. In Übereinstimmung mit Matsuda nehme ich an, daß die drei Pterygoten-Hauptgruppen sich früh in der Stammesgeschichte voneinander ge- trennt haben — im Gegensatz zu Matsuda ist die unterschiedliche Differenzierung ihrer Flug- apparate jedoch nicht auf einer Stufe anzuset- zen, auf der die Flügel noch als kleine Anlagen (“rudiments”) vorlagen, sondern später, als be- reits eine (wenn auch nicht sehr wirkungsvolle) Flugfähigkeit vorhanden war. Größere Verän- derungen geschahen wahrscheinlich erst nach der Aufspaltung der Pterygota in ihre Teilgrup- pen und gingen jeweils von einem ähnlichen (schon flugfähigen) Vorzustand aus. Demnach sind die drei verschiedenen Flugapparate der Pterygota — die keine eindeutigen Synapo- morphien aufweisen, die ein “Stück gemeinsa- mer Evolution” zweier Gruppen begründen könnten — zumindest in ihrem Grundplan als autapomorph anzusehen. Gegen eine unabhän- gige Entwicklung der Flügel der Odonaten und Ephemeropteren+Neopteren sprechen außer- dem auch die bis ins einzelne gehenden Über- einstimmungen zwischen den Ephemeropteren und Odonaten (etwa im dorsalen BAS-Bereich; vgl. S. 87f.), die Matsuda offensichtlich übersah. Die deutlichen Unterschiede zwischen den Flugapparaten sind andererseits nicht so er- staunlich, da in den drei Pterygoten-Linien drei verschiedene (alternative) Möglichkeiten von Antriebs- und Stellsystemen verwirklicht wur- den. Der Beitrag paläontologischer Forschung zu diesen Fragestellungen ist umstritten: 1) können fossile Flügel aus entsprechend weit zurücklie- genden erdgeschichtlichen Perioden in der Regel nicht sicher einer der rezenten Haupt- gruppen (oder ihrer Stammgruppe) zugeordnet werden — und 2) sind die für die Beurteilung des Flug-Funktionstyps wesentlichen proxima- len Bereiche der Flügel nur selten erhalten. In jüngerer Zeit wurden allerdings einige Ab- drücke beschrieben, die — nach den Abbildun- gen zu urteilen — zahlreiche Details der Gelenk- regionen erkennen lassen (Kukalova-Peck, 1974, 1978, 1983; Kukalova-Peck & Richard- son, 1983; Riek & Kukalovä-Peck, 1984); sie verdienen besondere Beachtung, v.a. weil sie evolutionstheoretisch sehr weitgehend interpre- tiert wurden, wodurch der Eindruck entstand, daß die große Lücke zu den Anfängen des In- sektenfluges nun endlich geschlossen werden könne. Die Flügelgelenkbereiche von Ostrava nigra und Mazonopterum wolfforum (Palaeodictyoptera der Familie Homoiopteridae aus dem Ober-Karbon, von Kukalova-Peck 1983 einer monophyletischen Gruppe “Paleo- ptera” zugeordnet) zeigen nach der Auffassung 108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 der Autorin die primitivsten Skleritanordnun- gen, die bisher bekannt sind. Sie führten zur Re- konstruktion einer aus 32 (!) Skelettelementen bestehenden ursprünglichen Gelenkregion der Pterygoten. Dieses “Ur-Flügelgelenk” wird sei- nerseits auf ein proximal der Subcoxa liegendes, zusätzliches Beinglied (“Epicoxa”) zurück- geführt, das seinen Exiten, den späteren Flügel, ringformig umgab (1983, l.c. Fig. 4). Nach der Zergliederung der Epicoxa in die 32 (zunächst gleichartigen, in Längs- und Querreihen an- geordneten) Teile seien dann die Gelenkstruk- turen der rezenten Pterygoten-Hauptlinien (Odonata+Ephemeroptera und Neoptera) durch unterschiedliche Kombination und Ver- schmelzung der Sklerite entstanden, wobei die Ur-Sklerite nach der Auffassung Kukalova- Peck’s bis ins einzelne gehend mit den Epipleu- riten, Axillaria etc. homologisierbar blieben (1983, l.c. Fig. 16). Die wenigen (und vagen) Aussagen zur Funktionsweise der Teile lassen die postulierten Entwicklungswege jedoch kei- neswegs “durchgängig” erscheinen. Es bleibt z.B. unklar, ob (und in welcher Ausprägung und Funktion) der Flügel seine Beweglichkeit überhaupt stets beibehielt, oder ob er nicht doch zeitweise eine Art unbewegliches “Para- notum” bildete (1983, l.c. S. 1634f.). Außerdem werden Muskulatur und Mechanik der rezenten Flugapparate zu wenig berücksichtigt. Wann und wie kamen die wesentlichen dorsalen Längsmuskeln (indirekten Senker), die das Ter- gum verwölben, in’s Spiel? Wie erklären sich die indirekten und direkten coxalen Flugmuskeln (die “bifunktionellen” Muskeln Wilson’s, 1962), die doch auf schon an der Pterygoten-Basis vor- handene, Insekten-typische Laufbeine schließen lassen und damit ein stabiles Pleurum-Widerla- ger erforderten? Kukalova-Peck muß (notge- drungen) bei ihren Aussagen zur Evolution der rezenten Konstruktionen sehr unbestimmt blei- ben und sich auf die allerersten Anfänge der Flügelentwicklung beschränken — die große Kluft zu den rezenten Apparaten bleibt offen. Ihre Hypothese beinhaltet im Grunde die An- nahme einer di-phyletischen Evolution der Flugfähigkeit (1983, l.c. S. 1638, 1645) — und ähnelt darin der Hypothese von Matsuda (1981), nur mit einer anderen Kombination der Gruppen!). Gerade weil aber ein funktioneller Brückenschlag zu den rezenten Gruppen fehlt, 1) Auch Matsuda (1981) lehnt übrigens eine mono- phyletische Entstehung der Flügel (als Stummel) nicht kategorisch ab! sind die vorgenommenen Sklerit-Abgrenzungen und “Homologisierungen” (1983, Fig. 16) zwei- felhaft und eigentlich beliebig; sie sind — in Er- mangelung wesentlicher Anhaltspunkte für die Homologisierung (z.B. der Muskulatur) — kaum zu belegen. Selbst bei den rezenten Insek- ten (die sich bis in winzigste Details studieren lassen) bieten die Gelenksklerite erhebliche Ho- mologisierungs-Schwierigkeiten; eine genaue Abgrenzung ist (infolge sekundärer Abgliede- rungen, Zerteilungen und Verschmelzungen) oft sehr schwierig, oder vorerst noch gar nicht möglich. Die durch “Nahtlinien” angedeuteten Grenzen und Schein-Grenzen der äußeren Morphologie lassen ja keine sicheren Rückschlüsse auf die Lage von Gelenkstellen und damit auf funktionelle Bewegungseinheiten zu (das gilt natürlich insbesondere für Fossi- lien). Gelenke können außerdem durch eine versteckte, z.B. versenkte Lage leicht übersehen - werden; v.a. aber liegen über die (für das Ver- standnis besonders wesentlichen) Gelenke der Flügel-Unterseite bisher keine Fossildokumen- tationen vor. In Anbetracht dieser Sachlage er- scheint es sehr wesentlich, daß bei der Beschrei- bung fossiler Flügel-Gelenkregionen äußerst vorsichtig vorgegangen wird und nur die wirk- lich eindeutig erkennbaren Teile dokumentiert werden. Da sowohl die Bildung monophyleti- scher fossiler Gruppen als auch ihre Zuordnung zu rezenten Einheiten auf große Schwierigkei- ten stößt, ist zudem eine begriffliche Klarheit unbedingt erforderlich (darauf geht Hennig, 1969, v.a. ım Teil I, in dem er die Möglichkeiten und Grenzen der Paläontologie analysiert, ausführlich ein). Die Widerlegung der von Hen- nig (1969) angeführten synapomorphen Merk- male der *Palaeoptera durch Kukalova-Peck (1983, S. 1661) scheint mir aus diesem Grund nicht fundiert zu sein, da *-Gruppen und (ech- te) Stammgruppen von der Autorin nicht klar unterschieden werden (ubrigens auch nicht bei Riek & Kukalova-Peck, 1984). Verschiedene Annahmen und Folgerungen Kukalovä-Peck’s lassen eine vorsichtig-kritische Einstellung etwas vermissen. Zu der (anschei- nend nicht in ihr Bild passenden) Tegula be- merkt die Autorin z.B. (1983, S. 1636): “The te- gula is a trichobothrium, not a sclerite (E. L. Smith, personal communication)...”. Nicht ein- mal bei den rezenten Gruppen ist jedoch eine zweifelsfreie Homologisierung der “Tegulae” möglich (vgl. S. 89)! Oder: Gleitflieger, die am Vorhandensein einer “starren” (hinteren) Axil- larplatte (= RAP) zu erkennen seien, “never Prau: Flugapparat der Libellen 109 flap their gliding planes while gliding” (1983, S. 1665). Zeigen segelnde Aeshniden (Aeshna grandis, Anax imperator) nicht sogar das Ge- genteil, nämlich alle Übergänge zum Schlag- flug? Außerdem besitzen auch die sicher höchst selten segelnden, wahrscheinlich plesiomor- phen (!) Zygopteren eine Radioanalplatte, die (wie bei Anısopteren) jedoch keineswegs starr ist. Eine genauere Zuordnung und Bewertung der fossilen Flügelgelenke (von denen man sich so viel versprochen hat) wird auch in Zukunft mit großen Schwierigkeiten verbunden sein. Ih- re Interpretation wird z.B. entscheidend vom Kenntnisstand der Funktionsmorphologie der rezenten Gruppen und von der Existenz einer Ausgangsbasis gesicherter Homologievorstel- lungen abhängen. Dies kann am Beispiel von Ostrava nigra noch einmal illustriert werden. Hennig vermutete (1969, l.c. S. 145; unter Be- zug auf eine frühere Arbeit Kukalova’s), daß das Flügelgelenk aller rezenten Pterygota “aus einem Vorzustand, wie ihn Kukalova bei Ostra- va beschreibt, ..., hervorgegangen sein muß”. Er ordnet Ostrava damit sogar weiter basal im System ein als Kukalová. Hennig empfindet allerdings das Fehlen der Costalplatte bei die- sem Fossil als etwas verwunderlich, wobei er davon ausgeht (lic. Abb. 25), daf} die ganze Odonaten-Costalplatte (incl. der dhCP) der Humeralplatte der übrigen Pterygoten homolog ist (in der vorliegenden Arbeit wird dagegen nur die dhCP der Humeralplatte gleichgesetzt). Neuere Abbildungen von Ostrava (Kukalova- Peck & Richardson, 1983, Fig. 20) zeigen im vorderen Basisbereich des Flügels einen weite- ren Skleriten, der in der Abb. 25 von Hennig noch fehlt. Dieser könnte (s. Kap. 3) mehrere Deutungen erfahren: Er könnte z.B. einer Hu- meralplatte (dhCP) entsprechen (die proximale Costalplatte, bzw. der Sklerit BAS, würde dann in einem basalen, nicht fossil erhaltenen Ab- schnitt liegen — sie könnte auch mehr oder we- niger reduziert sein), oder der proximal davon liegenden Struktur BAS homolog sein. Der ursprünglich an der Pterygotenbasis vorhande- ne Hauptteil der Costalplatte (BAS) wäre also bei Ostrava entweder noch vorhanden oder schon teilweise bis ganz reduziert. Die Hu- meralplatte könnte sogar auf die Flügeluntersei- te gewandert sein (vgl. S. 89f.). Eine genaue Zuordnung des (in wesentlichen Bereichen nicht erhaltenen) Flügelgelenks zu einer be- stimmten Gruppe (oder Stammgruppe) der Pte- rygota bleibt daher weiterhin problematisch. Evolution des Flugapparates innerhalb der Odonaten und funktionelle Deutung einiger Merkmale des Flügelgeäders Die Zygoptera und Anisozygoptera (mit der einzigen, durch zwei Arten vertretenen rezen- ten Gattung Epiophlebia) zeigen einige gemein- same Merkmale des Flugapparates, die auf eine ähnliche Flug-Spezialisierung (d.h. einen ahnli- chen Flugtyp) hinweisen. So ist der Winkel A der Grundschlagbahnebene des Flügels kleiner als bei den Anisoptera; er kann (beim Vorder- und Hinterflügel!) zur Vertikalen oder zur Ho- rızontalen hin verändert, also vergrößert oder verkleinert werden. Da die Anisozygoptera + Anisoptera eine monophyletische Gruppe bilden (beide Gruppen besitzen einige abgeleite- te, wahrscheinlich synapomorphe, Merkmale: Kiemenenddarm der Larven, s. Asahina, 1954; dreiarmige abdominale Haltezange der Männ- chen, s. Hennig, 1969, l.c. S. 321ff.), sind diese Ubereinstimmungen der Zygoptera und Aniso- zygoptera als Symplesiomorphien anzusehen (die Möglichkeit einer Konvergenz wird ausge- klammert, s. unten). Dafür sprechen auch Merkmale im Flugapparat der Anisoptera (wel- che selbst eine sichere monophyletische Gruppe darstellen — vgl. z.B. Fraser, 1957 und Pfau, 1971), die deutliche Anzeichen sekundärer Ver- änderung aufweisen: als Folge der Reduktion verschiedener Muskeln des Schlagbahn-Stellsy- stems — der pa fehlt in beiden Segmenten, der dim wurde im Metathorax reduziert (vgl. S. 61f.) — zeigen die Flugsegmente der Anısopte- ren insgesamt eine stärker ausgeprägte Hetero- nomie. Da es unwahrscheinlich ist, daß die ho- monome Ausprägung des Thorax bei Zygopte- ren und Anisozygopteren konvergent entstanden ist, kann gefolgert werden, daß der Pterothorax des letzten gemeinsamen Vorfahren der Odonata zygopteroid-anisozygopteroid be- schaffen war (vgl. dazu auch S. 82ff.). Interessant ist, daß bestimmte Teilstrukturen des Vor-Zurückschwingsystems innerhalb der Zygoptera (deren Monophylie nicht gesichert ist; vgl. Fraser, 1957) und bei Epiophlebia in un- terschiedlicher Ausprägung auftreten: Während im Mesothorax bei allen Gruppen zwei Tergal- sklerite (VTS und hTS) vorhanden sind, fand sich im Metathorax bei den meisten untersuch- ten Zygopterenfamilien (Calopterygidae, Epal- lagidae, Chlorocyphidae, Platycnemididae, Coenagrionidae und Protoneuridae) nur ein Sklerit, der vergrößerte vTS — der hTS ist redu- ziert, d.h. durch Membran ersetzt. Bei den Les- 110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 tiden und auch bei den Hemiphlebiiden!) ist der hTS dagegen auch im Metathorax noch weitge- hend erhalten, was als ursprünglicher Zustand aufgefafit werden kann. Epiophlebia besitzt im Metathorax ebenfalls beide Teile, gegeneinander noch beweglich, jedoch schon weitgehend aniso- pteroid verschmolzen (s.S. 62). Es erscheint lohnend, den Vergleich dieser Strukturen — die zwar funktionell relativ unbedeutend erschei- nen, aber auf eine monophyletische Teilgruppe der “Zygoptera” hinweisen (die einer Aufspal- tung der Zygopteren, wie sie Fraser, 1957, vorschlug, widerspricht!) — auf alle Gruppen auszudehnen. Aufgrund der mehr horizontal ausgerichteten Grundschlagbahnebene, und der Möglichkeit zur Schlagbahnveränderung bei Vorder- und Hinterflügel durch je zwei Stellmuskeln, sind die Zygopteren (und wohl auch die Anisozy- gopteren) auf engem Raum außerordentlich manövrierfähige Flieger, die in der Lage sind, auch den dichteren Bewuchs des Uferbereichs von Gewässern zu besiedeln (der primitive, noch sehr feindanfällige Kopulationsmechanis- mus ist in diesem Lebensraum anscheinend noch “tragbar”; vgl. Pfau, 1971). Die struktu- rellen und funktionellen Abwandlungen des Flugapparates in der Stammgruppe der Aniso- ptera — die Verstellung der Grundschlagbahn- ebene und die Einengung des Schlagbahn-Stell- bereichs der Flügel — verändern diesen ursprünglichen Flugtyp wesentlich. Sie können als eine Anpassung an eine neue Ökologische Zone interpretiert werden: Die rezenten Aniso- pteren stellen robustere, schnellere Flieger dar, die auch besser in der Lage sind, Dauer- und Streckenflüge zu vollbringen. Sie haben den Luftraum über der freien Wasserfläche erobert. Infolge der vertikaler stehenden Grundschlag- bahnebene ihrer Flügel wird (beim Auf- und Abschlag) mehr Vortrieb erzeugt, was den be- sonders “reißenden” Flug erklärt. Anscheinend wurden verschiedene Muskeln (die “Rück- schwing”-Muskeln pa im Mesothorax und dlm im Metathorax; vgl. S. 61f.) mit der Verstellung der Schlagbahnebene in der Stammgruppe der Anisoptera überflüssig und konnten reduziert 1) Hemiphlebia mirabilis Selys (die einzige rezente Hemiphlebiide) besitzt im Meso- und Metathorax außerordentlich ähnlich ausgebildete Sklerite vTS und hTS — wahrscheinlich ein sehr ursprünglicher Zustand. Für die Überlassung eines Exemplars von Hemiphlebia sei Herrn Dr. J. A. L. Watson an die- ser Stelle herzlich gedankt. werden. Damit ging auch eine Vereinfachung der Flugmechanik einher, was wiederum den Flügelantrieb effizienter machte. Im Metathorax wurde zusätzlich der Flügel-Vorschwingmecha- nismus (und sein Muskel, der pa) reduziert. Dieses Segment wurde so im Zuge der Speziali- sierung der Anisoptera zu “Vortriebsfliegern” zu einem weitgehend reinen Antriebssegment (die tergale Mechanik konnte in diesem Zusam- menhang besonders stark vereinfacht werden! Vgl. S. 62). Die Veränderung der Grund- schlagbahnebene beider Flügel bedingte jedoch gleichzeitig einen Verlust an Auftrieb, der an- scheinend dadurch aufgefangen werden konnte, daß die Flügelspreiten — v.a. im Hinterflügel, in dem der Analteil besonders vergrößert ist — verbreitert wurden. Man könnte erwarten, daß der pronatorisch- supinatorische Drehspielraum der Flügel bei den Anisoptera mit der Einschränkung des Schlagbahnspielraums verkleinert werden konn- te. Bestimmte, gegenüber den Zygopteren abge- wandelte Merkmale des Flügelgeäders deuten jedoch sogar eher auf eine erweiterte Verwin- dungsfähigkeit der Flügel hin; sie stehen wahr- scheinlich im Zusammenhang mit der Verbrei- terung der Flügel oder/und der Verstärkung des Flügelantriebs. (Da diese Geäder-Veränderun- gen v.a. den Aufschlagsdrehbereich betreffen, erweitern sie möglicherweise in erster Linie den Spielraum der Vortriebserzeugung; s. oben und S. 93ff.) So ist bei den Anisopteren das Flügel- dreieck (die Discoidalzelle) sekundär ver- größert, zweigeteilt und durch zusätzliche Adern stabilisiert (vgl. Fraser, 1957). Dadurch wird ein größerer distaler und kaudaler Flügel- bereich an die Cubitalsektor-Basis angekoppelt, wodurch sowohl die Ubertragung der aktiven Kräfte nach distal (auf die größere Flügelfläche) wie auch der passiven Kräfte nach proximal (zur Flügelbasis) gesichert oder verbessert wird. Wahrscheinlich ist außerdem die Verlagerung des Ursprungsorts der Flügeladern “IR,” und “Ry,5” (Abb. 8) zur Flügelbasıs hin (vgl. Fraser, 1957) in einem Zusammenhang mit der Verwin- dungsfähigkeit größerer (breiterer) Flügel zu se- hen, da auch dadurch die von diesen Adern “ge- tragenen” Flügelbereiche enger an den Arculus und die Cubitalsektor-Basis angeschlossen wer- den (vgl. auch Pfau, 1975). Dieses Merkmal tritt allerdings auch bei bestimmten Teilgruppen der Zygoptera (und bei den Anisozygoptera) auf, und zwar v.a. dort, wo breitere, weniger “ge- stielte” und dichter geaderte (schwerere!) Flügel vorhanden sind. Fraser (1957) stellt diese Grup- Prau: Flugapparat der Libellen 111 pen in die Nahe der Anisoptera (+Anisozygo- ptera) und betrachtet die Zygoptera daher als uneinheitliche (paraphyletische) Gruppe. Wenn sich diese Hypothese erharten ließe, hatte man gleichzeitig ein gewichtiges Argument dafür, daß der gestielte, schmale (“typische”) Zygo- pterenflügel als der ursprüngliche Flügel der Odonata (*Odonata! Vgl. Fußnote S. 76f.) an- zusehen ist. Der nähere Anschluß von “IR,” und “R,,;” an den Arculus (und die dadurch er- reichte Erhaltung oder Verbesserung der Ver- windungsfähigkeit im Aufschlagsdrehbereich) könnte bei Zygopteren jedoch auch konvergent entstanden sein. Dafür spricht z.B., daß inner- halb verschiedener Zygopterenfamilien noch Übergänge der Aderverlegung erhalten sind. In einigen Gruppen (Epallagidae, Calopterygidae) ist die Merkmalsevolution sogar weiter voran- geschritten als bei den Anisoptera: “R,,,” ent- springt dort beinahe direkt vom Arculus. In einem funktionellen Zusammenhang mit der Pronations-Supinations-Drehmechanik des Flügels stehen sicher noch zahlreiche weitere Merkmale des Flügelgeäders. Die Stabilisierung des Costalsektors durch verstärkte Costa-Ra- dius-Queradern (“antenodal primaries”, vgl. Fraser, 1957; pan, und pan,, Abb. 3) spielt z.B. . im Aufschlagsdrehbereich eine wichtige Rolle, da der Cubitalsektor ein solides Verwindungs- Widerlager benötigt. In den breiteren Flügeln der Anisoptera+Anisozygoptera (und einiger Gruppen der Zygoptera!) ist der Costalsektor durch weitere Queradern zusätzlich verstärkt. Auffallend ist dabei, daß der Arculus am Co- stalsektor meist in der Mitte zwischen den bei- den (durch ihre besondere Dicke hervorgehobe- nen) “primaries” artikuliert — manchmal ist die Gelenkstelle des Arculus auch der ersten oder zweiten Hauptquerader genahert, die dann be- sonders verstarkt ist. Die genauere Kenntnis der Flügelmechanik ermöglicht jetzt sogar die funk- tionelle Interpretation einiger ganz unbedeu- tend erscheinender Merkmale des Odonaten- Flügelgeäders: Der schräge Verlauf des Arculus (Abb. 3 und 8) ist z.B. wohl v.a. der “Distal- Komponente” der supinatorischen Bewegung des CuS (s.S. 53) “angepaßt” — eine entgegen- gesetzte Schrägstellung des Arculus würde die Verwindung behindern. Die Ausrichtung der kurzen, ebenfalls besonders kräftigen (aber in den Abbildungen von Odonatenflügeln meist nicht berücksichtigten) proximalen Costa-Ra- dius-Querader cr, (etwa senkrecht zur Achse El, s. Abb. 3 und 7) steht dagegen in einem Zu- sammenhang mit dem Flügeldreh- und Verwin- dungsmechanismus des Abschlagsdrehbereichs; auch hier würde eine andere Ausrichtung der Ader (und eine andere Lage des Gelenkpunkts c3, an dem die Ader vorn endet) die Verwin- dungsbewegung beeinträchtigen. Weitere vergleichend-morphologische wie auch biomechanische Untersuchungen sind für eine bessere Beurteilung der Flügelgeäder- Merkmale notwendig. Sie könnten nicht nur un- ser Verständnis der Funktionsweise der Flügel der rezenten Pterygoten (und ihrer mannigfalti- gen Funktionswechsel) erweitern, sondern wür- den gleichzeitig auch Einblicke in die Biologie der ausgestorbenen Gruppen vermitteln, von denen wir, als einzige Reste, oft nur Flügel- geäderteile kennen. Dre MUSKELFUNKTIONEN IM TABELLARISCHEN VERGLEICH Die Komplexitàt der thorakalen und pteralen Mechanik macht es in vielen Fallen schwer, Muskelfunktionen durch einfache Zugexperi- mente an den Sehnen auf Anhieb zu erkennen; die zahlreichen Irrtumsmöglichkeiten können erst bei genauer Kenntnis aller Einzelmechanis- men und ihrer wechselseitigen Abhängigkeit re- duziert werden. Entsprechend vielfältig ist das Bild, das sich beim Studium der Literatur ergibt. Selbst die Autoren, die sich eingehender mit dem Flugapparat der Libellen beschäftigten, Tannert (1958) und Russenberger & Russenber- ger (1959/60), weichen in ihren Ergebnissen zur Skelettmechanik — und in den davon ausgehen- den Interpretationen der Muskelfunktion — stark voneinander ab. Neville (1960) unternahm den Versuch, die Wirkungen der Muskeln durch Beobachtung von Skelettbewegungen und durch Muskelausschaltexperimente bei vor dem Windkanal fliegenden Libellen direkt zu erken- nen. Dabei ergaben sich jedoch offenbar Wi- dersprüche, die (ohne genauere Kenntnis der Skelettmechanik) nicht aufzulösen waren. Außerdem ist es Neville wohl nicht gelungen, die Bewegungen der Skelett-Teile tatsächlich mefßbar zu machen. Da seine Meßpunkte in Wirklichkeit sehr komplizierte Raumbahnen beschreiben (die Bewegungen stehen in Abhän- gigkeit von mehreren Kräften und können sich zusätzlich überlagern), erfordert die Erfassung der Bewegung eines Punktes (in Bezug auf eine Kraft) nicht nur perspektivische Korrekturver- fahren, sondern die gleichzeitige Berücksichti- gung aller anderen wirkenden Kräfte. Es ist da- her nicht gerechtfertigt, dat Neville seine Er- gebnisse als exakt (weil “quantitativ”) von 112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 “nur” qualitativen Befunden positiv abhebt. In der Tabelle 1 sind die Muskel-Termini und -Funktionen verschiedener Autoren zusammen- gefaßt. Dabei mußten manche Funktionsbe- zeichnungen in den hier verwandten Begriffsge- brauch “übersetzt” werden. In den anschlie- ßend unter 1. bis 23. angefügten Bemerkungen zur Tabelle wird versucht, einige abweichende Ergebnisse zu erläutern und bis zu ihren theore- tischen Voraussetzungen (die manchmal allerdings schwer ersichtlich waren) zurückzu- verfolgen. 1. — Clark (1940) geht nicht näher auf die Skelett- mechanik ein, so daß die Muskelfunktionen z.T. nicht erklärt sind. Die Basalarmuskeln (dvm; ,) werden den “Subalarmuskeln” (pm, _,) als antagonistische Mus- keln der Flügeldrehbewegungen gegenübergestellt: bei zeitlich verschiedenem Einsatz (“alternate action”) würde entweder der vordere Teil des Flügels (Prona- tion) oder der hintere (Supination) gesenkt; gleichzei- tige Aktion beider Muskelgruppen würde dagegen zur Senkung des ganzen Flügels führen (l.c. S. 556). Die mit diesen Angaben nur angedeutete (einfache) Flügeldrehmechanik konnte nicht bestätigt werden. Tabelle 1 CLARK, 1940 TANNERT, 1958 RUSSENBERGER und "RUSSENBERGER, 1959/60 dvm, Heber Hebermuskel (2-geteilt) 2) dvm Heber, Supinator-Promotor | (bzw. Pronator-Remotor) | dvm; Senker 1) vorderer Senkermuskel 2) bm1 Senker, Pronator (-Remotor) | Pronator Pronator | pm, Senker 1) mittlerer Senkermuskel À sml Senker (Supinator-Promotor) | Supinator | dvm, Heber — — | dvm, Heber? — — Beinbeweger? dvm, Senker 1) Steuermuskel Costa bm2 ‘ Pronator Pronator, Senker ©) 2) | pm, Pronator Steuerm. Ventralausläufer Pt1 Tpm (1,2) | (Mesothorax Anisoptera), Pronator *) | Seitl. Verspannerm. d. Mesoscutum (Zygoptera) u. Metascutum (Zygoptera + Anisoptera) dvm, Heber Steuermuskel Lateralauslaufer Pt1 Supinator 5) | axm Remotor (Pronator) dvm, Heber Steuermuskel r + m — c ) Heber, Supinator pm, Senker 1) hinterer Senkermuskel 2) sm2 Supinator (Promotor) !°) | Supinator (etc. s. l.c. S. 411, 422) pm,, | 1) Steuermuskel Analis 1a, (Senker) Pronator 6) pm; | Supinator Steuermuskel Analis 1b, dim Supmaton (rene ton) Supinator 6) pm; Senker 1) Steuermuskel Analis 2 DI) sim Supinator (Promotor) Supinator Senker (etc. s. l.c. S. 411, 422) dim, Senker hinterer Verspanner dim Pronator-Remotor 2) des Mesoscutum 8) (bzw. Supinator-Promotor) = = smi Remotor (Pronator) 123) vgl. S.112— 117 ‘VILLE, 1960 it tergosternal ber, Pronator Prau: Flugapparat der Libellen 113 2. — Costal- und Radioanalplatte werden von Tan- nert (1958) als funktionell scharf getrennte Flügelbe- reiche betrachtet (s. zum Beispiel lc. S. 423). Die Flügelverwindung kann daher nach Tannert durch alle diejenigen Muskeln beeinflußt werden, welche eine Basisplatte, relativ zur anderen, auf- oder abwärts be- wegen können. Tannert nımmt dies für fast alle Mus- keln an, auch für die Haupt-Heber und -Senker (l.c. S. 424; für den bas1, und für den als antagonistisch ange- sehenen hca, wird die Verwindungsfunktion z.B. auf S. 419 näher ausgeführt). Dabei wird zwischen einer Hauptfunktion (Antrieb) und einer Nebenfunktion (Verwindung) unterschieden. Es wird jedoch nicht HATCH, 1966 =) dvm1 Heber naher erlautert, wie sich die zahlreichen, und zudem auf ganz verschiedene Weise (vgl. auch Anmerkung 3 und 4) den Flügel verwindenden Einflüsse distal im Flügel auswirken und gegenseitig beeinflussen. Auch Hatch (1966) nımmt offensichtlich eine unab- hangige Auf- und Abschlagsbewegung der CP relativ zur RAP (und umgekehrt) als Ursache der Pronation und Supination an (l.c. S. 709). Hatch und Tannert kommen jedoch z.T. zu entgegengesetzten Ergebnis- sen. Ein (distales) Senken der Costa durch den bas1 ergibt bei Tannert eine pronatorische Drehung, wel- cher der hca (durch Heben der Costa) entgegenarbei- ten könne (l.c. S. 410, 419); bei Hatch sind diese Mus- PFAU, 1986 dvml Aufschlagsm. (Antrieb) (Ruckschwingm.) ond basalar 15) 16) 18) dvm3 Senker, Pronator 20,22) bas 1 Abschlagsm. (Antrieb) iker, Pronator tsubalar D) pml Senker 20) subl Abschlagsm. (Antrieb), Supinator dvm2 21) dvm2 Einstellm. (Abschlag) dvm5 21) (tc; s. Fußnote S. 45) t basalar 15) 16) 18) dvm4 Pronator 20) 21) bas2 Einstellm. (Aufschlag) iker, Pronator | pm5 Promotor (?) tp Einstellm. (“Klickmech.”) erior coxoalar 7) dvm6 Heber, Supinator 20) Nev ca Supinator der unteren dinator, Remotor, Schlagwende (Heber) ber sterior coxoalar dvm7 Heber, Pronator 20) hea Pronator der oberen ber (l.c. S. 639) Schlagwende (Heber) ond subalar NS) pm2 Senker, Supinator BNP) sub 2 Supinator (Abschlag), iker, Supinator Senker pm4a,b (Funktion s.b. Tannert) fa Pronator (Aufschlag) rd subalar 15) 19) pm3 Supinator 20) 20) subs Supinator + Einstellm. iker, Supinator (Aufschlag) dlm1 Promotor (?) 23) dlm Vorschwingm. (Mesothorax) Rückschwingm. (Metathorax) a pa ny Rückschwingm. (Mesothorax) Vorschwingm. (Metathorax) 114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 keln dagegen (beziiglich der Fligeldrehung) Syner- gisten (zu Hatch vgl. auch Anm. 20—23). 3. — Tannert’s Bewegungsanalysen sind anschei- nend durchweg auf mazeriertes Tiermaterial gegrün- det. Beim Mazerieren wird das Resilin aus den Ge- lenkverbindungen (wie z.B. dem Pleuralgelenk pi) herausgelöst; diese zeigen anschließend einen viel weiteren Bewegungs-(Membran-)Spielraum als beim lebenden oder frischtoten Tier. Das Pleuralgelenk p1 (“Unterstützungsgelenk der proximalen Cp” bei Tan- nert) wurde sicher aus diesem Grund als “Schlittenge- lenk” verkannt (l.c. S. 404, 418, 420; Abb. 12, 13). Damit wohl zusammenhangend ist nach Tannert die gesamte CP (bei Tannert: proximale Cp+distale Cp; hier, etwas verandert: vCP+mCP+phCP+dhCP, vgl. S. 41) um eine Achse drehbar, die vom “vorde- ren Scuto-Alargelenk” (t1) durch das vordere Pleural- gelenk (p1) zum “Bereich des distalen Verwindungs- gelenks” (bei c3) läuft (l.c. Abb. 5, 18, 36; der distale “Angelpunkt” dieser Achse wird dabei offensichtlich vom Gelenk c3 unterschieden — l.c. S. 418, 423, 432 — ein zweites Gelenk konnte von mir jedoch an die- ser Stelle nicht aufgefunden werden). Diese Verwin- dungsachse existiert jedoch in Wirklichkeit nicht; eine Drehung der gesamten Costalplatte (relativ zur RAP+“c-RAP-Brücke”, l.c. S. 418) kann nicht statt- finden (zur Achsen-Anordnung und -Funktion vgl. auch Abb. 7 und S. 47ff.). Der bas2 ist damit kein Drehmuskel der CP (Pronator); bas2 und vca (die sich bei Tannert — an einem zwei-armigen Hebel angrei- fend — zu beiden Seiten der “Verwindungsachse” ge- genüberstehen) sind außerdem nicht als Antagonisten zu betrachten (vgl. auch Anm. 4). 4. — Nach Tannert ist der vordere Tergalsklerit vTS (den er, zusammen mit dem Randsklerit, als “Pte- rale 1” bezeichnet — zur Homologisierung des Ptera- le 1 vgl. jedoch S. 83f.) um eine in Tierlängsrichtung verlaufende Drehachse schwenkbar (l.c. Abb. 14b). Der Tergopleuralmuskel tp (“Steuerm. Ventralausläu- fer Pt1”) soll nun durch Anhebung des Randsklerits (RS — bei Tannert “Lateralausläufer Pt1”) indirekt (durch Schub von ventral her auf die CP) eine prona- torische Drehung der gesamten CP (vgl. Anm. 3) be- wirken; der Muskel ist bei Tannert somit ein Antago- nist des vca und ein Synergist des bas2. Eine Bewe- gung des vTS durch den tp nach lateral-ventral (zur Anhebung des Randsklerits; vgl. l.c. Abb. 14b) ist je- doch nicht möglich. Der RS ist außerdem als ein-ar- miger, am vTS gelenkig ansıtzender Hebel zu betrach- ten, über den bei Kontraktion des vca — durch Zug an der hinteren Costalplatte hCP — eine Supination erreicht wird; der tp wird dabei nicht gedehnt. Vorde- rer Tergalsklerit und Randsklerit bilden demnach kei- nen zwei-armigen Hebel, wie Tannert annımmt. Eine Schubwirkung des wenig harten (biegbaren) RS kann — auch aufgrund der Zwischenmembran zur phCP — ausgeschlossen werden. 5. — Die genaue Muskelwirkung wurde nicht er- läutert (zu der nach Tannert abweichenden Funktion des mesothorakalen tp der Anisoptera s. Anm. 4). Nach der vorliegenden Arbeit haben die tp-Muskeln im Meso- und Metathorax (bei Zygopteren und Anisozygopteren+ Anısopteren) die gleiche Funktion (s. S. 45f.). 6. — Die beiden Muskeln arbeiten nach Tannert an- tagonistisch und heben bzw. senken das “Analfeld” (l.c. Abb. 28). Die “Verwindungsachse” würde dabei proximal durch die Mitte des Ansatzgebietes der bei- den Muskeln, distal mitten durch den Arculus verlau- fen (l.c. Abb. 5, 36). Diese Ausrichtung der Drehachse und die antagonistische Funktion zweier Muskelteile des fa (von den beiden Seiten eines zwei-armigen He- bels aus) konnte nicht bestätigt werden (vgl. S. 53ff.); das “Analfeld” Tannert’s entspricht daher auch nicht dem Cubitalsektor. Eine Zweiteilung des fa wird (wie im Falle des dvm1, vgl. Fußnote S. 60) durch die ein- tretende Trachee, und eine dafür vorhandene Ein- buchtung im Apodem, vorgetäuscht. 7. — Tannert nimmt anscheinend eine “Steuerung” der großen Hauptmuskeln durch die kleinen (als syn- ergistisch interpretierten) Nebenmuskeln an (lc. S. 424f.) — sowohl bezüglich der Schlagfunktion als auch der (für fast alle Muskeln postulierten) Verwin- dungsfunktion (s. Anm. 2). Vor allem für die Paare bas1—bas2 und sub2—sub3 wird dies genauer erklärt (s. z.B. l.c. S. 411). Nach der hier dargestellten Auf- fassung (S. 45) “steuern” die kleinen, tonisch aktiven Muskeln dagegen die Kraftwirkung des jeweils anta- gonistischen Hauptmuskels (der bas2 also z.B. den dvm1; für den sub3 vgl. S. 56). 8. — Der Muskel wird bei Tannert nur auf S. 409 (für den Mesothorax) ohne nähere Erklärung seiner Funktion erwähnt. 9. — Russenberger & Russenberger (1959/60) neh- men eine kombinierte Flügel-Pronation und -Remo- tion durch die Muskeln sm3, axm und dim beim Abschlag an; entgegengesetzt (supinierend und vor- .ziehend) wirken beim Aufschlag der dvm (s. aber wei- ter unten) und auch der dım (zur indirekten Wirkung des dım s. auch Anm. 13). Diesen Flügelbewegungen wird eine komplizierte Mechanik (u.a. Kippung des Tergum “in der Medianebene”, d.h. um eine Quer- achse, und Annäherung der pleuralen “Schwingen- pfeiler”) zugrundegelegt (l.c. Abb. 31a,c). Fast alle Muskeln (auch die direkten Antriebs-Senker) spielen, dieser Mechanik zufolge, eine Rolle als Pronatoren- Remotoren bzw. Supinatoren-Promotoren (einige Funktionen wurden in der Tabelle — der von Russen- berger & Russenberger postulierten Mechanik ent- sprechend — in Klammern ergänzt). Die beiden Autoren betrachten den dvm (ähnlich wie Tannert) als 2-köpfigen Doppelmuskel, wobei sie im dorsalen An- Prau: Flugapparat der Libellen 115 satzapodem ein Quergelenk (zwischen den beiden Muskelteilen) annehmen (vgl. lc. S. 23f., 48). Bei Kontraktion würde der dvm entweder Pronation-Re- motion oder Supination-Promotion bewirken, je nachdem welcher Teil des Muskels stärker arbeitet. Die in Abb. 31a, c bei Russenberger & Russenberger dargelegte Mechanik konnte nicht bestätigt werden (zur Zweiteilung des dvm vgl. Fußnote S. 60). 10. — Russenberger & Russenberger diskutieren für diese Muskeln, die nach ihrer Ansıcht kaum als Senker von Bedeutung sein können, die Möglichkeit einer “Rücksteuerung” des Thorax-Resonanzsystems (vgl. Lc. S. 80ff.). 11. — Die Autoren erwägen eine rechts-linksseitig unterschiedliche Kontraktion dieser Muskeln und nehmen (sehr allgemein) eine Beeinflussung der “Stel- lung der einzelnen, am Flugmechanismus beteiligten Sklerite” (lc. S. 28) an. Sie diskutieren auch die Mög- lichkeit einer gegenläufigen Bewegung des rechten und linken Flügels, u.a. bewirkt durch die Tpm (l.c. S. 47 und Abb. 31b; eine ähnliche Funktion der Tpm= pm5 nimmt auch Hatch, l.c. S. 713, an). Der dafür als Beweis angesehene Hochschulfilm von v. Holst, 1950, zeigt diese Gegenlaufigkeit jedoch nicht (vgl. dazu S. 43f. und S. 45f.). Obwohl Russenberger & Russen- berger auf die doppelfrequente Bewegung der Pleuren hinweisen (l.c. S. 46f.), erkannten sie die Bedeutung der Tpm für die Einstellung eines bistabilen Flü- gelschlagmechanismus anscheinend nicht. 12. — Russenberger & Russenberger untersuchten offensichtlich die Muskulatur subadulter (aus Larven gezogener) Exemplare von Aeshna cyanea (l.c. Abb. 18). Darauf weisen die Muskelproportionen und auch der (im adulten Tier reduzierte) Muskel sm3 (s. auch Anm. 14) hin. Einige Fehleinschätzungen sind wohl darauf zurückzuführen: der sm3 wird anscheinend mit dem Illısm, von Clark (1940) verwechselt (einem zum Abdomenrand führenden, intersegmentalen Muskel, der entgegen Clark’s Annahme ohne Bedeu- tung für den Flügelschlag ist), den die beiden Autoren nun auch im Mesothorax aufzufinden meinen; vca und hca (=axm) werden nicht unterschieden (s. l.c. S. 28), der Ursprungsort außerdem nicht richtig be- schrieben (Verwechslung mit anderen Muskeln?); der von Russenberger & Russenberger als dlm bezeichne- te Muskel (fa der vorliegenden Arbeit) wird im Meta- thorax anscheinend mit dem (dort bei juvenilen Tieren noch größeren) dım vermengt und als ein Muskel be- trachtet; etc. 13. — Russenberger & Russenberger beschreiben eine (auf dem Schlagphasen-Unterschied zwischen Vorder- und Hinterflügel beruhende) unterschiedli- che, d.h. pronatorische (beim Abschlag) oder supina- torische (beim Aufschlag) Drehwirkung dieses Mus- kels in den beiden Schlagphasen (lc. S. 77f.; s. dazu auch Anm. 9). Sie nehmen außerdem an, daß der Mus- kel dım die “Phasenverschiebung zwischen Vorder- und Hinterflügel” beeinflußt (l.c. S. 26, 49: “Koppel- muskel zwischen den Flügelsegmenten”). Die seg- mentkoppelnde Kraft der dım ist jedoch infolge der starken Spreizung der Muskeln nach kaudal wahr- scheinlich sehr gering (vgl. S. 61). 14. — Dieser Muskel, von Russenberger & Russen- berger als “3. Subalarmuskel” bezeichnet, ist wohl mit dem Pleuroalarmuskel pa (dem Muskel Nr. 31 bei Asahina, 1954 — im Metathorax Nr. 53) identisch. Er läuft bei Zygopteren (und bei der Gattung Epiophle- bia) vom kaudalen Innenrand der RAP (oder der Membran medial davon) nach vorn-seitlich-unten an die Pleuralleiste (Abb. 2). Bei adulten Anisopteren ist der Muskel reduziert (nur eine Sehneneinstülpung zeigt noch seine dorsale Ansatzstelle) — bei subadul- ten kann er dagegen in beiden Segmenten noch mehr oder weniger gut entwickelt aufgefunden werden (vgl. dazu Anm. 12). Asahına (1954) beschreibt den Muskel auch für Davidius (möglicherweise untersuchte er ein subadultes Exemplar); ich konnte ihn jedoch bei an- deren (adulten) Gomphiden (/ctinogomphus, Onycho- gomphus, Gomphus) nicht mehr auffinden. 15. — Zur Erklärung der Drehwirkung der Flügel- senker und -heber bezieht sich Neville (1960, S. 631, 653) auf eine Hypothese von Weis-Fogh (1956). Da- nach stehen sich bei Schistocerca im Mesothorax vor und hinter dem Fulcrum Antriebsmuskeln gegenüber (die Basalar- und Subalarmuskeln), die als Antagoni- sten (durch verschieden starken Einsatz oder unter- schiedlichen Kontraktionszeitpunkt — s. auch Anm. 1 zu Clark, 1940) die Flugelanstellung, durch Dre- hung des Flügels um das Fulcrum, bestimmen sollen. Der sub1 wird von Neville dementsprechend — auf- grund seiner angenommenen “neutralen Lage” zwi- schen den vor dem Fulcrum befindlichen Basalarmus- keln (die als Pronatoren angesehen werden) und den dahinter liegenden Subalarmuskeln sub2 und sub3 (welche den Flügel supinieren) — als reiner Ab- schlagsmuskel betrachtet (l.c. S. 649). Nach der in der vorliegenden Arbeit beschriebenen Verwindungsme- chanik trifft das jedoch nicht zu (vgl. S. 47ff.). Auch bei Heuschrecken hat sich gezeigt, daß der Flügel nicht einfach als Ganzes um das Fulcrum drehbar ist .(— diese Bewegung würde mit dem Auf-Abschlags- mechanismus des TWM in Konflikt geraten), sondern daß eine die Antriebsmechanik “umgehende”, flü- gelinterne Verwindungsmechanik vorliegt (vgl. Pfau, 1977b, 1978a; s. dazu auch S. 91, 92). 16. — Unglücklicherweise wird der 1. Basalarmus- kel (first basalar bzw. dvm, bei Clark) von Neville als “second basalar”, der 2. Basalarmuskel (second bas- alar Clark’s oder dvm,) dagegen als “first basalar” be- zeichnet. Beide Basalarmuskeln werden als phasische Muskeln betrachtet, die schon am Ende des Auf- schlags (l.c. S. 649; evtl. sehr frühzeitig, s. Fig. 19f — vgl. aber Anm. 18) eingesetzt werden können. Zum Nachweis ihrer pronatorischen Wirkung (die in der vorliegenden Arbeit nicht bestätigt werden kann; vgl. 116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 S. 50) durchtrennte Neville beide Basalarmuskeln und beobachtete die Flügelanstellung bei vor dem Wind- kanal fliegenden Tieren. Da die Flügelschlagfrequenz nach Ausschaltung um etwa ein Drittel zurückging, ist die beobachtete veränderte Flügelanstellung (lc. Fig. 19e) jedoch für eine Pronationsfunktion der Bas- alarmuskeln nicht unbedingt beweiskräftig: mit einer geringeren Flügelgeschwindigkeit wird ja gleichzeitig die passive Pronation verringert bzw. ist auch eine Zunahme der supinierenden Wirkung der Muskeln sub1 und sub2 zu erwarten (ganz abgesehen von der Möglichkeit, daß die Libelle bei Kappung von Mus- keln den Einsatz anderer Muskeln zum Ausgleich ver- ändert hat). . 17. — Der Muskel kann nach Neville seine Supina- tions-Wirkung schon vor dem Abschlagsende (l.c. Fig. 13 und v.a. Fig. 21) und dann auch in der 1. Hälf- te und fast ganzen 2. Hälfte der Aufschlagsphase (l.c. S. 643, 653, 655) entfalten. (Neville’s Fig. 9 und 13 stimmen jedoch damit und untereinander nicht ganz überein.) Bei der Erklärung der vermuteten zusatzli- chen Funktion des vca als Remotor (l.c. S. 653) treten Widersprüche auf: wenn eine Flügel-Vorbewegung (nach Neville durch den vca, gegen Ende des Ab- schlags) mit einem abrupten Auseinanderweichen von RAP und CP einhergeht (l.c. Fig. 11d, 14, 15; S. 640f.; vgl. dazu aber S. 99f. der vorliegenden Arbeit), dann kann der vca kurz danach, an der Schlagwende, nicht auf einmal die entgegengesetzte Bewegung verursa- chen und zu einem Remotormuskel werden. Das er- neute Schließen des Spaltes zwischen den beiden Flügelbasisplatten (l.c. Fig. 11e) wird von Neville an- scheinend damit erklärt, daß der vca, nach einer nur anfänglich starken Kontraktion, sich (plötzlich) nur noch schwach weiter kontrahiert (l.c. S. 655) und da- durch (und durch seine Aufschlagsfunktion? s. unten) eine Flügel-Remotion bewirkt. Neville nımmt außerdem eine “automatische” Rückziehwirkung der Aufschlagsmuskeln, und eine entsprechende, passive Vorziehwirkung der Ab- schlagsmuskeln, an (lc. S. 652f., 641), wobei er jedoch nicht zwischen der Schlagbewegung in der (festliegen- den) Grundschlagbahnebene und der (davon weitge- hend unabhängig möglichen) Vor-Zurückschwing- Bewegung unterscheidet. Zur Erklärung einiger Phänomene mußte Neville dann zu komplizierten Zu- satzhypothesen greifen (lc. S. 652f.). Der Autor beobachtete eine Diskrepanz zwischen natürlichen und manipulierten Flügelschlägen (lc. S. 641f., 653) und zog diese z.T. zur Erklärung der vca- Funktion heran. Die beobachteten Unterschiede kön- nen jedoch anders gedeutet werden. Sie beruhen wohl darauf, daß Neville den Vorderflügel bei seiner künst- lichen Imitierung des Schlags supiniert angestellt ab- wärts bewegte: der supiniert angestellte Vorderflügel schwingt am Abschlagsende (ausweichend) nach vorn (und “supiniert” dabei weiter, vgl. S. 59) und dann am Beginn des Aufschlags sofort wieder zurück, wo- bei sich RAP und CP abrupt nähern (entsprechend Neville’s Fig. 14, 15). Dieses Vor- und Zurück- schwingen ist jedoch keinesfalls die zwangsläufige Folge der geneigten Flügelschlagbahnebene (lc. S. 641), sondern ergibt sich (zufällig) aus der unnatürli- chen Flügelmanipulation und der (nur am Abschlags- ende und Aufschlagsbeginn vorhandenen) Flügelbe- weglichkeit nach vorn und zurück. Daß sich der Randsklerit (der Ansatzzipfel des vca, bei Neville “anterior lobe of the lateroprescutum”) bei einem ma- nipulierten Flügelschlag im Unterschied zum natirli- chen Schlag nicht abrupt nach ventral bewegt, ist an- dererseits nicht verwunderlich. 18. — Der sub2 kann nach Neville supinierend in die Auf-Abschlagswende eingreifen (l.c. Fig. 19f.; S. 649) und dabei, und dann v.a. beim Abschlag (Fig. 19e), die pronierende Wirkung der Basalarmuskeln kontrollieren. Der Autor übersah jedoch andere pro- nierende Kräfte (die pronierende Wirkung des hca; die beim Abschlag passiv-pronierende Luft); die Pro- nationsfunktion der Basalarmuskeln trifft nach der vorliegenden Arbeit andererseits nicht zu. Die von Neville aus den Experimenten abgeleitete antagonisti- sche Beziehung bas1,2—sub2,3 kann somit nicht be- statigt werden. Ein fruher Einsatz der Muskeln sub2 und bas1 (“second basalar” bei Neville) an der Auf- Abschlagswende ist zwar wahrscheinlich möglich, wurde jedoch durch das Ausschaltexperiment (lc. Fig. 19f) nicht erwiesen (zur Beweiskraft der Basalar- muskel-Durchtrennungen fur die von Neville postu- lierte pronatorische Funktion der Muskeln vgl. Anm. 16). Der 2. Subalarmuskel wird in der vorliegenden Ar- beit als der wesentliche Gegenspieler der passiv-pro- nierenden Luft in der Abschlagsphase betrachtet (s.S. 50f., 93ff.), der basl ist dagegen als reiner Senker anzusehen (s.S. 50). 19. — Nach Weis-Fogh (unpubliziert, s. Neville, Lc. S 648) ist dieser Muskel tonisch aktiv (vgl. dazu auch S. 44). Neville beschreibt mehrere Funktionen: 1) eine (kombinierte) Abschlags- und Supinations- funktion innerhalb der Abschlagsphase (l.c. S. 648, 654) sowie 2) eine Supinationsfunktion (zeitlich nach der Supination durch den vca) in der zweiten Hälfte der Aufschlagsphase (s. l.c. Fig. 19 und S. 647, 655). Bedenkt man jedoch die hier dargelegte zugfederähn- liche Wirkung tonischer, mit der Antriebsmechanik in Beziehung stehender Muskeln, so wird der sub3 beim Abschlag entlastet und ist somit in dieser Phase “aus- geschaltet” (vgl. S. 56). 20. — Den Text erklärende Funktionsskizzen feh- len leider. Aus der Beschreibung (v.a. l.c. S. 709, 713) geht hervor, daß Hatch (1966) das Zustandekommen der Pronations-Supinations-Bewegungen auf ver- schiedene Weise erklärt: a) Durch proximales Heben bzw. Senken der CP und RAP, die (voneinander unabhängig) um ihre pleuralen Auflagepunkte schwenken und den distal der Gelenkpunkte liegen- den, dazugehörigen Flügelteil entweder senken oder Prau: Flugapparat der Libellen 17, heben. Senkt sich z.B. die CP proximal, wird der Flü- gel supiniert, senkt sich die RAP proximal, dann wird er proniert. Diese Mechanik erinnert an Tannert (s. Anm. 2). b) Durch Kippung des Tergum um eine Querachse: senkt sich das Tergum-Vorderende, wirkt dies nach Hatch auf die CP und ergibt (s. a)) eine Su- pination; senkt sich das Hinterende, wirkt dies auf die RAP und hat (s. a)) eine Pronation des Flügels zur Folge. Diese Annahme einer tergalen Beweglichkeit um eine Querachse erinnert an Russenberger & Rus- senberger, führt aber zu gerade entgegengesetzten Flügeldrehungen (vgl. auch Anm. 9, sowie Abb. 31c bei Russenberger & Russenberger). c) Durch prona- torische oder supinatorische Drehungen der RAP, die proximal mit entsprechenden Verkippungen des Ter- gum (etc. s. b) und a)) einhergehen. Die pronatorische Wirkung des dvm7 (hca) wird nach Hatch anscheinend auf zweierlei Weise erreicht: durch (distales) Heben und durch Supinieren (!) der RAP (lc. S. 713); die letztere Bewegung verstellt dann wohl das Tergum so, daß die CP proximal angeho- ben wird, was nach a) ebenfalls eine Pronation ergibt (? — oder es handelt sich hier um einen Druckfehler, s. auch weiter unten). Da der dvm7 die RAP jedoch eindeutig pronatorisch dreht, kann Hatch’s Mechanik schon aus diesem Grund nicht zutreffen. Auch der bei Hatch beschriebene Supinations-Mechanismus des dvm6 (vca) — durch distales Heben der CP (s. a)) — kann in Wirklichkeit nicht stattfinden: da CP und RAP über zwei distal der Schlagachse liegende Ge- lenkpunkte miteinander verbunden sınd (vgl. S. 50), würde durch ein distales Anheben der CP der ganze Flügel nur aufgeschlagen. Ausgehend von der postulierten CP-RAP-Unab- hangigkeit, nach der Unterschiede in der normalen Schlagbewegung der beiden Basisteile zu Änderungen der Flügelanstellung führen, betrachtet Hatch die vor- deren direkten Senker (dvm3 und dvm4) als Pronato- ren, die hinteren (pm2 und pm3) als Supinatoren (die letzteren sollen jedoch auch die RAP supinieren kön- nen, was im Widerspruch zum oben beschriebenen dvm7-Mechanismus steht). Der pmi (lc. Fig 6A) wird in dieser Hinsicht als neutral betrachtet (was aber anderen Aussagen widerspricht; s. a)). 21. — Für die als tonısch eingeschätzten Muskeln dvm2, dvm4, dvm5 (?) und pm3 nimmt Hatch an, daß sie den Flügel beim Gleitflug in ihrer Stellung (und z.T. auch in der Anstellung) stabilisieren. Sie sollen außerdem für die Flügel-Ruhehaltung bedeutsam sein. 22. — Der pm2 steht dem dvm3 nach Hatch beim Abschlag antagonistisch gegenüber (vgl. dazu auch Anm. 18 zu Neville). 23. — Hatch nimmt außerdem an, daß die (ebenfalls als tonisch eingeschätzten) mesothorakalen dlm1 die Vorder- und Hinterflügel beim Flug voneinander ent- fernt halten. DANKSAGUNG Die Untersuchungen wurden durch die Deut- sche Forschungsgemeinschaft finanziert (Ri 57/18, Rı 57/20-3 und Pf 174/1-1). Für ihre Ermöglichung und geduldige Förderung danke ich Herrn Prof. Dr. H. Risler herzlich, für wertvolle Diskussionen und Durchsicht des Manuskripts v.a. Herrn Prof. Dr. D. Bilo (Saarbrücken) und Frau Dr. B. Schroeter. Zu besonderem Dank bin ich außerdem Herrn Dr. S. Asahina (der mir fixierte Exemplare der außergewöhnlichen Libelle Epiophlebia super- stes überließ), Herrn Prof. Dr. G. Rüppell (der mir eine Kopie seines hervorragenden Filmes über das Flugverhalten von Aeshna cyanea zur Auswertung zusandte) und Frau K. Rehbinder (die verschiedene Abbildungen uberaus sorgfal- tig “ins Reine” zeichnete) verpflichtet. ZUSAMMENFASSUNG Der Libellenflügel wird beim Schlag in einem (durch zwei pleurale Gelenklager gebildeten) Scharniergelenk bewegt — dadurch ist eine Grund-Schlagbahnebene festgelegt. Drei starke Muskeln treiben den Flügel an, die direkten Senker bas1 und subl und der indirekte Heber dvm1. Schwächere, tonische Muskeln (dvm2; bas2, sub3) können entweder den Abschlags- oder den Aufschlagsmuskeln entgegenwirken, so daß der “Flugmotor” in beiden Schlagphasen (auch unilateral) “gedrosselt” werden kann. Der Muskel tp vermag eine elastische, bistabile Komponente des Flügelschlags — durch Verän- derung der Rückstellkraft der Pleuralleiste, die in beiden Schlagphasen zunächst nach lateral ausgelenkt wird — einzustellen. Die Drehungen des Flügels um die Längsach- se (Pronation, Supination) laufen in zwei me- chanisch unterschiedlichen Drehbereichen, die den beiden Schlagphasen zugeordnet werden können, ab. Bei einer Pronation im Abschlags- drehbereich wird der “Verstellflügel” sowohl als Ganzes (in zwei proximalen Scharniergelen- ken) proniert als auch gleichzeitig unter Span- nung gesetzt und pronatorisch verwunden; die Verwindung wird dabei durch die Schubbewe- gung eines vorderen Flügelteils (Costalsektor) gegen den dahinter liegenden Hauptteil der Flügelspreite (Cubitalsektor) bewirkt. Bei einer Supination im Aufschlagsdrehbereich bewegt sich dagegen der Cubitalsektor gegen den Co- 118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 stalsektor. Der Flügel wird dadurch ebenfalls verwunden. Diese Verwindung ist jedoch (im Gegensatz zur pronatorischen Verwindung im Abschlagsdrehbereich) eine “reine” Flügelver- windung — sie geht mit keiner Drehung des ganzen Verstellflügels einher. Die beiden Dreh- bereiche grenzen in einer mittleren Anstellung (0°) aneinander; sie sind dort durch mechani- sche Anschläge (die ein “Uberlappen” verhin- dern) voneinander getrennt. Die verschiedenen pronatorischen bzw. supinatorischen Stellmus- keln sind somit entweder dem einen oder dem anderen Drehbereich zuzuordnen. Einzelne dieser Muskeln können als Muskeln der Schlag- wendepunktsdrehungen angesehen werden (hca; vca), andere als Einstellmuskeln der Flügelanstellung. Unter den letzteren ver- größern bestimmte Muskeln den aerodynami- schen Anstellwinkel und arbeiten dabei gegen die anströmende Luft (subl und v.a. sub2 beim Abschlag, fa beim Aufschlag); nur ein Muskel, der sub3, vermag den Anstellwinkel zu verklei- nern und verringert gleichzeitig die Aufschlags- geschwindigkeit. Die Kontraktion der mesothorakalen dorsa- len Längsmuskeln (dlm) bewirkt eine Schubbe- wegung des Tergum nach kaudal-dorsal und führt — in einem flügelinternen, aus zwei Ge- lenken zusammengesetzten Scharniergelenk — zu einem Vor-Schwingen des Flügels (Ande- rung der Schlagbahn). Dieser Bewegungsablauf kann jedoch erst gegen Ende des Abschlags, und nach Erreichen des 0°-Anschlags des Abschlagsdrehbereichs, stattfinden. Der dvm1 (der Heber des Flügelantriebs) schwingt den Flügel zu Beginn des Aufschlags (automatisch) wieder zurück. Die bei Zygopteren und Anıso- zygopteren (Epiophlebia) gegenüber den An- isopteren (v.a. im Metathorax) abweichenden Verhältnisse des “Vor-Zurückschwing-Sy- stems” werden beschrieben und diskutiert. Die dargelegten Ergebnisse zur Skelettme- chanik unterscheiden sich von älteren Befunden in wesentlichen Punkten und führen dement- sprechend zu neuen Interpretationen der Flug- und Stellmuskel-Funktionen. Die Hypothesen der verschiedenen Autoren werden verglichen. Bestimmte Mechanorezeptoren in der Flügel- basıs — ein Chordotonalorgan (CH) und zwei Reihen von campaniformen Sensillen (CF1, CF2) — werden durch die Verwindungsbewe- gungen des Flügels beansprucht: das CH wird sowohl bei pronatorischer als auch supinatori- scher Verwindung über kleine Hebelelemente der Flügelunterseite entdehnt (gedehnt wird der Rezeptor jeweils bei den Rückdrehungen zur 0°-Anstellung hin); in der Kutikula der Felder campaniformer Sensillen treten bei den Fligel- verwindungen Zugspannungen (quer zum Flügel) auf. Elektrophysiologische Ableitungen (bei Drehung des Flügels um die Längsachse) ergeben, daß die Sensillen des CH stark pha- sisch sind und Flügeldrehungen in beiden Dreh- bereichen (und dort in beiden Richtungen) an- zeigen. Die campaniformen Sensillen, die ein phasisch-tonisches Zeitverhalten aufweisen, werden dagegen nur bei Drehbewegungen zu den Anstellextremen hin erregt — CF1 anschei- nend bei Pronation, CF2 bei Supination. Diese Rezeptoren könnten zur Messung der geometri- schen Flügelanstellung und (indirekt — über den jeweiligen Spannungszustand in der Kuti- kula und dessen zeitliche Anderung) zur Regi- : strierung der die Flügelanstellung verändernden Luft- und Muskelkräfte eingesetzt werden. Der Vergleich der Flugapparate der Odona- ten, Ephemeropteren und Neopteren führt zu neuen Homologievorstellungen und zur Rekon- struktion eines “Ur-Flugapparates” der Ptery- goten (mit zwei Schlagachsen); die rezenten Flugapparate können davon ausgehend in drei alternativen Linien abgeleitet werden. Damit kann der Hypothese einer polyphyletischen Entstehung der Pterygota widersprochen wer- den — das Problem der Aufspaltung der Ptery- gota bleibt jedoch ungeklärt. Evolutive Verän- derungen innerhalb der Odonaten betreffen v.a. die Ausrichtung der Grundschlagbahnebene und das Vor-Zurückschwing-System (das nur bei Zygopteren und Anisozygopteren in beiden Pterothorax-Segmenten entwickelt ist); die Anısoptera werden als sekundär vereinfachte “Vortriebsflieger” angesehen. SUMMARY During the wing stroke the dragonfly wing moves up and down on a hinge joint formed by two pleurum-to-wing articulations, which de- termines the stroke plane angle of the wing. The wing is driven by strong direct depressor mus- cles (bas1, sub1) and indirect levators (dvm1), which provide motor power for flight. Certain tonic muscles, which are comparatively weak (dvm2, bas2, sub3), are able to counteract either the downstroke (dvm2) or the upstroke (bas2, sub3) power muscles; thus the flight motor can be throttled back in both stroke phases. A ter- Prau: Flugapparat der Libellen 119 gopleural muscle (tp) is able to vary the recoil of the pleural ridge, which is bent outwards and loaded in the first part of both the downstroke and the upstroke, and swings back inwards (and is unloaded) in the second part. In this way a bistable mechanism, superimposed on each wingstroke phase, is adjustable by the muscle tp. The mechanics of the wing movements along its long axis (pronation, supination) are diffe- rent in two ranges of rotation (“Abschlagsdreh- bereich”, “Aufschlagsdrehbereich”), which pre- sumably correspond to the ranges of geometri- cal angle of attack used within the two stroke phases. During pronation in the “Abschlags- drehbereich” a major part of the wing (“Verstellflügel”) is pronated as a whole. Since this movement is determined by two proximal hinge joints, the wing is additionally put under pressure and also pronated by twisting; the twisting is caused by the pressure of the frontal sector of the wing (“Costalsektor”) against the caudal sector (“Cubitalsektor”). During supina- tion in the “Aufschlagsdrehbereich” it is the cu- bital sector, which is pressed against the costal sector; this again causes a twisting (supination- twisting in this case), but is not associated with a movement of the “Verstellflügel” as in the “Abschlagsdrehbereich”. The two different ran- ges of rotation border on each other at 0°, the mean geometrical angle of attack, at which the wing is not twisted. Mechanical stops prevent overlapping of the ranges of rotation. Therefore the muscles of pronation and supination can be assigned to either the “Abschlagsdrehbereich” or the “Aufschlagsdrehbereich”. Different types of pronator and supinator muscles are described: (a) muscles that rotate the wing at the turning-points of the stroke (up- per turning-point: hca; lower one: vca) and (b) muscles that are able to adjust the angle of at- tack mainly within the upstroke or downstroke phase. Among these latter muscles some exert force against the airflow (rotating the wing into the opposite direction), increasing the aerody- namical angle of attack either in the upstroke (fa) or downstroke (subl, sub2) phase. Only one muscle (sub3) is able to reduce the angle of attack during the upstroke — at the same time reducing the speed of the wing (see above). Contraction of the dorsal longitudinal mus- cles (dlm) causes the tergum to shift in a caudal and dorsal direction, resulting in a forward swinging of the wing; most important for this movement is a hinge joint consisting of two single joints both lying at the base of the wing, For mechanical reasons the resulting alteration of the stroke plane angle can only occur at the, end of the downstroke — after the 0°-stop of the “Abschlagsdrehbereich” has been reached. The muscle dvm1 (main levator of the flight motor) is able to swing the wing backwards (automatically) at the beginning of the upstroke. Zygoptera and Anısozygoptera (Epiophlebia) differ from the Anisoptera in some details (mainly ın the metathorax) of this “Vor-Zu- rückschwing-System” of the wings. In particular they possess direct antagonistic muscles (dlm— pa) in both segments of the pterothorax (in this regard they are considered plesiomorphous). Surprisingly, these muscles show opposite func- tions in the mesothorax and the metathorax. Previous studies of sceletal mechanics show widely differing results (compared to one an- other and to the present study). Various conclu- sions of these studies concerning the functions of musculature are compared and discussed. Certain mechanoreceptors, lying in the base of the wing — a chordotonal organ (CH) and two rows of campaniform sensilla (CF1, CF2) — are mechanically stressed during the prona- tion and supination movements. For example, during pronation-twisting as well as during su- pination-twisting (beginning at 0°) CH is shor- tened via two small levers on the underside of the wing, which are in contact with the costal sector or cubital sector respectively. The recep- tor is stretched during opposite movements, i.e. during supination in the “Abschlagsdrehbe- reich” and pronation in the “Aufschlagsdrehbe- reich”, and reaches its maximum length at the 0°-stops. In the cuticular zone of the campani- form sensilla (CF1 and CF2) tension stresses (transverse to the long axis of the wing) increase when the wing is twisted (pronated or supi- nated) and decrease as the geometrical angle of attack falls to 0°. Electrophysiological investiga- tions reveal a strongly phasic response of the CH in both directions of movement for both ranges of rotation. The campaniform sensilla however, which are phasic-tonic, only spike if the wing is twisted. CF1 presumably records the pronation-twisting and CF2 the supination- twisting. These latter receptors could therefore measure the geometrical angle of attack and also indirectly, via the specific patterns and courses of cuticular tensions, provide information con- 120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 cerning the aerodynamical and muscular forces that affect the angle of attack of the wing. A comparison of the flight apparatus of Odo- nata, Ephemeroptera and Neoptera reveals pos- sible new homologies and allows of the recon- struction of an ancient flight system of the Pte- rygota (“Ur-Flugapparat”), which possessed two main axes of wing stroke. The modern forms of the flight apparatus can be derived from this in three functionally differing lines of evolution, which are exclusive of one another. This contradicts the hypothesis of a polyphyle- tic origin of the Pterygota. However, the pro- blem of the phylogenetic splitting of the Ptery- gota into three main groups remains unsolved. Within the group *Odonata, evolutionary de- velopments have mainly concerned the stroke plane angle and the “Vor-Zurückschwing- System”. The latter is present in both mesotho- rax and metathorax only in the Zygoptera and Anisozygoptera. it is almost entirely lacking in the metathorax of the Anisoptera, which are considered as specialized for forward-thrust flight. cl (C1) c2 (C2) c3 c4 el, 2 (E1, 2) Ef Prau: Flugapparat der Libellen ABKURZUNGEN Analis hca vorderes pleurales Flügelschlag-Gelenk (p1 hCH bei Odonaten) 1. Flügel-Schlagachse (durch a und b hCP bestimmt; P1/P2 bei Odonaten) HP Antecosta des Metatergum hTS Arculus (Querader R+ M/CuP) kMB Gelenk des Arculus am Radius M 2. Flügel-Schlagachse (verläuft durch b) m “Abkömmlinge” der Achse B bei Odonaten (entsprechen P2/C4 und C2/C4) mCP hinteres pleurales Flügelschlag-Gelenk (p2 MP1 bei Odonaten) ms Basis-Sklerit (s.S. 79f.) n Teile des BAS bei Ephemeropteren Pmax Basalarsklerite der Neopteren Basalarmuskeln pl Sinnesborstenfelder Costa p2 proximales Gelenk der HP (c2 bei Odonaten) P1/P2 Gelenk 1 in der CP (zwischen mCP und phCP) — Großbuchstabe kennzeichnet die P2/C4 Drehachse Gelenk 2 in der CP (zwischen phCP und pa dhCP) — Großbuchstabe kennzeichnet die pan, 2 Drehachse phCP Gelenk 3 der CP (zwischen dhCP und CoSB PN bzw. RAP) Pel Gelenk 4 der CP (zwischen dem mCP- Ptl 2 Kaudalfortsatz und der RAP: Flügelunterseite) TRE durch die Gelenke c2 und c4 laufende R Scharnierachse der Vor- r Zurickschwingbewegung RAP Felder campaniformer Sensillen RS Chordotonalorgan Cor Costalsektor Costalsektor-Basis Sc Costalplatte “ScH? Costalplatte ohne dhCP Costa-Radius-Queradern sub1-3 Cubitus posterior ij Cubitalsektor tl Cubitalsektor-Basis Cubitalsektor-Hebel t2 distales Gelenk der HP (c3 bei Odonaten) distale hintere Costalplatte T1/T1 dorsaler Langsmuskel Tb Dorsoventralmuskeln tp kaudales Gelenk der Ventralseite des BAS “TPM+TWM” nach distal (in den Flügel) versetztes Gelenk e bei Odonaten (entspricht c4) TPM nach ventral (ins Pleurum) versetztes Gelenk e bei Neopteren TWM1 Scharniergelenke des Epifulcrum — grofe Buchstaben kennzeichnen die Drehachsen TWM2 Epifulcrum Fulcrum (hinterer pleuraler Gelenkkopf) TZ ventrales Gelenk des bas I der Neopteren a (Kap. 3) vat Falz in der RAP-Oberseite (nicht Kap. 3) Fulcroalarmuskel CP basales Biegegelenk des Fulcrum vGK 1. und 2. Gelenksklerit der RAP vTS Gelenk zwischen Tb und T 3 Hebelapodem der Tergalbrücke 121 hinterer Coxoalarmuskel hinterer Hebelsklerit des Chordotonalorgans hintere Costalplatte Humeralplatte (dhCP bei Odonaten) hinterer Tergalsklerit kaudale Media-Basis Media Membranzone in der (medialen) Wand des dvm1-Apodems mittlere Costalplatte Mittelplatte 1 der Neopteren Membranspalt am CuSH Nodus-Gelenk extrem pronierte Anstellung (Abschlagsdrehbereich) vorderes pleurales Flügelschlag-Gelenk, zwischen vGK und CP (mCP) hinteres pleurales Flügelschlag-Gelenk, zwischen Fulcrum und RAP (Ef) Schlagachse des Flügels (verläuft durch pl und p2) 2. Hauptachse des Abschlagsdrehbereichs (verläuft durch p2 und c4) Pleuroalarmuskel primäre Antenodal-Queradern (“primaries”) proximale hintere Costalplatte Postnotum Pterale 1 möglicherweise (gemeinsam) dem Pti homologe Sklerite bei Ephemeropteren “Pterale 4” der Ephemeropteren (vgl. S. 85) Radius Resilin Radioanalplatte Randsklerit extrem supinierte Anstellung (Aufschlagsdrehbereich) Subcosta “Scutellarhebel” der Ephemeropteren (vgl. S. 85) Subalarmuskeln mittlere Tergalregion vorderes Tergalgelenk des Flügels (zwischen Tb und vCP) hinteres Tergalgelenk des Flügels (zwischen TZ und RAP) Scharnierachse der Tb-Bewegung Tergalbrücke Tergopleuralmuskel “Ur-Antriebsmechanismus” der Flügel (vgl. S. 78ff.) Tergalplatten-Mechanismus, Flügelantrieb der Odonaten Tergalwölbungs-Mechanismus (1), Flügelantrieb der Ephemeropteren Tergalwolbungs-Mechanismus (2), Flügelantrieb der Neopteren Tergalzapfen vorderer Coxoalarmuskel vorderer Hebelsklerit des Chordotonalorgans vordere Costalplatte vorderer (pleuraler) Gelenkkopf vorderer Tergalsklerit Zone verstärkter Kutikula in der kaudalen RAP 122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 3, 1986 LITERATUR Asahina, S., 1954. 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Biol. 91: 1—24. NT a apici) 7 wee ion Steet ay er deer es £3 ét is VE N . ne an ti Mi (al N 1 i ! LA ì MA HA ti y ae DEEL 129 AFLEVERING 4 1986 DL 46 | T 563 Et be TH DSCHRIET VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING \ I, INHOUD J. C. Roskam. — Biosystematics of insects living in female birch catkins. IV. Egg- larval parasitoids of the genera Platygaster Latreille and Metaclisis Forster (Hymenoptera, Platygastridae), pp. 125—140, figs. 1—44. Tijdschrift voor Entomologie, deel 129, afl. 4 Gepubliceerd 10-XI-1986 LA vi En i = hi , 5 ry. pr | a È D à > eo BIOSYSTEMATICS OF INSECTS LIVING IN FEMALE BIRCH CATKINS. IV. EGG-LARVAL PARASITOIDS OF THE GENERA PLATYGASTER LATREILLE AND METACLISIS FORSTER (HYMENOPTERA, PLATYGASTRIDAE)... J.C. ROSKAM by FA f à Division of Population Biology, University of Leiden, The er ABSTRACT Adult and larval stages of Platygaster betularia Kieffer, P. betulae (Kieffer) and ER sis phragmitis Debauche are described! These species are egg-larval parasitoids of three gall midge species, which belong to the genus Semudobia Kieffer (Diptera, Cecidomyiidae), in fruit catkins of Betula (Betulaceae). The various developmental stages of the Platygaster species are discriminated with the help of multivariate methods. Phenology, host specificity and effects upon host density have been investigated. All platygastrid parasitoids develop highly synchronized with their hosts. Platygaster betularia and P. betulae have mutually exclusive host preferences. Both Platygaster species are important mortality factors particu- larly able to eliminate moderate host densities. Speciation patterns in Semudobia and Platy- gaster have no parallel traits and can, therefore, not be regarded as results of a co-evolution- ary process. Some notes are included about platygastrid parasitoids of Nearctic Semudobia species and of inquiline Dasineura gall midges in birch catkins. INTRODUCTION Gall midges allied with female birch catkins are frequently attacked by parasitoids belonging to the hymenopterous superfamilies Scelionoi- dea and Chalcidoidea. The scelionoid represen- tatives are the object of this study. They are egg-larval endoparasitoids: the eggs are laid in the host egg, but further development does not occur before the host is in its final instar. Until then parasitized hosts can not be distinguished from healthy ones. Parasitized early final instar hosts become inert and further development eventually ceases. One, or sometimes two larvae are visible inside the host, consuming all of the host’s body contents within a few days. The skin of the host larva remains as a “cocoon”, providing an extra protection for the mature parasitoid larva, in which it pupates. Kieffer (1916) described two platygastrid B asitoids of Semudobia betulae (Winnertz) s.l viz., Platygaster betularia Kieffer and Misco) clops betulae Kieffer. According to current opinion, also adopted in this paper, Platygaster Latreille and Misocyclops Kieffer are synony- !) A formal synonymy will be proposed by Mr. H. J. Vlug, Wageningen (pers. comm.). mous, because male diagnostic characters do not allow a grouping of the involved species into two genera!). Fulmek’s (1968) compilation ex- cepted, later reports on egg-larval parasitoids of Semudobia mention only P. betularia (Barnes, 1951; Bachmaier, 1965). Hodges (1969) treated the life-history of this parasitoid. All these au- thors considered the gall midge fauna of female birch catkins as relatively simple: Semudobia betulae, the gall maker, is accompanied by a sa- prophagous and a predaceous gall midge spe- ‚cies, viz., Clinodiplosis cilicrus (Kieffer) and Lestodiplosis cf. vorax (Rubsaamen), respective- ly. Roskam (1977, 1979) and Roskam & van Uf- felen (1981), however, arrived at the conclusion that at least five gall inducing Semudobia species and two inquiline Dasineura species are special- ized on female birch catkins. Clinodiplosis cili- crus and Lestodiplosis cf. vorax are frequently present in this biocoenosis. The advancement of knowledge at the gall midge level provided a ba- sis for further research of the parasitoids and the results of this study are now presented for the egg-larval parasitoids. Platygaster betularia and P. betulae are both abundant in the Palaearctic entomofauna of fe- male birch catkins. Among other things they are 125 126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 b RÉAL LL LD Ss WILL ILL % ZZ m Sc eam In Muy È S = LU Ke S = \ Nun = À LENS \ > > LE sag SSL GEL. LTE WIE aa DI MLE SEES = = S S S SI Ÿ S Figs. 1—16. Adult platygastrid characters. 1, 13, fore and hind wing, female; 2, 5, antenna, female; 3, 4, 6, an- tenna, male; 7—9, ultimate and penultimate antennal segments, male; 10—12, ditto, female; 14—16, gaster, fe- male. 1, 5—7, 10, 16, Metaclisis phragmitis; 2, 3, 9, 12—14. Platygaster betularia; 4, 8, 11, 15, P. betulae. b, basal vein; m, medial vein; os, oval sensilla; s, stigma; sc, subcostal vein. 1, 13—16, x 60; 2—6, x 96; 7—12, x 240. characterized by different host associations: P. betularia has only been reared from S. betu- lae (Winnertz) s.s. and $. skuhravae Roskam, whereas P. betulae is restricted to S. tarda Ros- kam. Metaclisis phragmitis Debauche has been reared from both S. betulae and S. tarda. Also the Nearctic gall midge S. brevipalpis Roskam is attacked by a platygastrid: mature larvae have been found in one collection. These larvae are very aberrant in shape and belong to an undes- cribed genus. Dasineura species have other platygastrid parasitoids. Because mature Dasineura larvae drop to the ground for hibernation, it was not possible to rear parasitoid full grown larvae and adults from these inquilines. However, in Dasi- neura larvae two different forms of platygastrid larvae have been observed: the first putatively belongs to Piestopleura cf. mamertes (Walker), the second could not be combined with adult platygastrids frequenting female birch catkins. No platygastrid parasitoids have been found in Clinodiplosis and Lestodiplosis larvae. MATERIAL AND METHODS Immature stages. — Galls of different Semu- dobia species have different shapes and can therefore be sorted according to the gall induc- Roskam: Platygaster and Metaclisis 127 ing midge species (Roskam, 1977). In order to detect the parasitoids, the host larvae were dis- sected from identified galls and macerated in warm 80% lactic acid. Platygastrid larvae were taken from opened hosts and slide-mounted in polyvinyl-lactophenol. Galls were also collected from dry herbarium material. Then 10% KOH was used for maceration. Adults. — Adults were collected by rearing them from samples of identified galls, and by collecting ovipositing females from female cat- kins with an exhauster. This material was either stored in 80% ethanol, or mounted on tags, or dissected and slide-mounted in euparal. Speci- mens representing all stages of the studied spe- cies have been deposited in the collection of the Rijksmuseum van Natuurlijke Historie, Leiden. Phenological observations on immature stages were made by analyzing samples of ten fruit catkins each. The samples were collected weekly from the beginning of March until the end of September. Adults were caught from mid-April until the end of May. Every day, dur- ing a period of ca. 30 min. around noon, about twenty female wasps and a similar sample of gall midges were collected and subsequently identi- fied. Host-parasitoid specificity was determined by rearing adult parasitoids from gall samples sorted according to the gall maker. Mortality caused by parasitoids was defined by dissecting gall samples that had been collected in Decem- ber. All seasonal activities have then ended, but many fruit catkins are still complete and can be collected from the trees. An extensive description of the study-areas Meijendel (52.08N 4.20E), Duivenvoorde (52.06N 4.24E), Kootwijk (52.11N 5.46E), Il- perveld (52.29N 4.58E) and Nieuwkoop (52.10N 4.50E) was given in Roskam (1977); Hulshorst (52.22N 5.44E) and Kootwijk are dry areas on sand. ADULTS Adults of species belonging to Platygaster were described by Kieffer (1926) and those of Metaclisis phragmitis by Debauche (1947). Therefore, attention will be paid here only to some differential characters. Metaclisis. — (figs. 1, 5—7, 10, 16). Fore wing with subcostal and medial vein, basal vein indicated by a more or less distinct dark streak, subcostal vein terminated by a distinct stigma, which does not reach the front margin of the wing. Second (sex) flagellomere in male as wide as third, without large, oval sensilla. Proximal part of female second gastral tergite broad, about % as wide as distal part. Sheaths of ovi- positor exposed. Platygaster. — (figs. 2—4, 8, 9, 11—15). Wing venation reduced. Second (sex) flagello- mere of male wider than third, with large, oval sensilla. Proximal part of female gastral tergite about half the width of the distal part. P. betularia. — Males. Flagellomeres subqua- drate, length of fifth flagellomere less than 1.4 times its width in lateral view (fig. 3). Proximal part of scutellum rather dull, due to relatively rich setation (fig. 37). Females. Scutellum as in male (fig. 39). Gaster twice as long as wide, gradually narrowing to- wards its apex (fig. 14); exposed part of fifth and sixth segments about a fifth (0.19—0.22, n=5) the length of the gaster without oviposi- tor; surface of fifth segment shiny, without any sculpture. P. betulae. — Males. Flagellomeres oblong, length of fifth flagellomere more than 1.4 times its width in lateral view (fig. 4). Proximal part of scutellum shiny, due to relatively sparse setation (fig. 38). Females. Scutellum as in male (fig. 40). Gaster © © © ui length ultimate flagellomere (um) SJ © 320 360 400 440 width head (um) 480 Fig. 17. Species differences in Platygaster females. The ellipses indicate 95% confidence limits. Dots, P. betulae: asterisks, P. betularia. 128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 Table 1. Mean values (x) and standard deviations (s) of adult characters, measurements in um. MALES FEMALES betularia betularia b-lae/b-ria betulae betularia skuhravae betularia betulae overlap skuhravae 366.6 31.9 | 369.0 22.2 | 377. width head 42.9 |336.1 23.4 | 358.7 28.4 | 357.4 35.5 557.2 55.9 | 550.8 35.4 | 544. length gaster length ult. flagellomere length anh flagellomere h width at flagellomere more than twice as long as wide (5:2), distinctly narrowed between fourth and fifth segment (fig. 15); exposed part of fifth and sixth segments about 0.3 times (0.26—0.31, n=5) the length of the gaster without ovipositor; surface of fifth gastral segment with band of longitudinal striae. Means and standard deviations of five variates for different species of Platygaster, as well as for different host groups of P. betularia, are pre- sented in table 1. Interspecific percentages of overlap, or percentages of misclassification, are lowest for the length of the ultimate flagello- mere, as well in males, as in females. This char- acter provides therefore the best univariate dis- crimination of P. betularia and P. betulae. A more powerful interspecific discrimination is obtained by various combinations of character pairs: (width head — length ultimate flagello- mere), (width head — length fourth flagello- mere), (length gaster — length ultimate flagello- mere) and (length fourth flagellomere — width same segment) in males; (width head — length ultimate flagellomere) in females. All these com- binations provide amounts of misclassification below 1%. The latter combination is plotted in fig. 17. For explanation of the technique, univa- riate as well as bivariate, see Lubischew (1962). In order to obtain the best separation between the two Platygaster species, a multivariate func- tion, viz., discriminant analysis, was carried out. This technique has been explained by Pimentel (1979) and was applied by Roskam (1982). Per- centages of misclassification after application of discriminant analysis remained 0.02 in males and 0.17 in females. In table 2 the values of the character set are summarized. In males, the length of the fourth flagellomere contributes most to discrimination, whereas in females, as 47.4 1590.9 49.1 | 633.1 60.1 | 736.0 89.2 was expected on results of univariate analysis, the length of the ultimate flagellomere is most important for species discrimination: these . characters scored the highest values as coeffi- cients for the canonical variates, 0.5198 and 0.2914, respectively. The canonical score Z (sum of the products of character values and corresponding coefficients for canonical va- riates, table 2) is plotted in fig. 41. Identification of new specimens is possible by calculating their canonical score Z (Bigelow & Reimer, 1954; Roskam, 1982). Such identification runs then as follows: Zmales = (length fourth flagellomere) + 0.32 (length ultimate flagellomere) —0.21 (width head) < 4 belden AE dsl P. betularia P. betulae (length ultimate flagellomere) + 0.47 (length fifth flagellomere) — 0.68 (width fourth flagellomere) < N ae P. betularia Z females = Idem>84........... P. betulae P. betularia has been reared in considerable numbers from two different hosts, Semudobia skuhravae and S. betulae. No discrimination between specimens reared from different hosts was possible in an univariate way (table 1). Also in a multivariate way, viz., discriminant analy- sis, no discrimination was possible between subgroups of P. betularia which developed in different host species: in males misclassification remained 40.5%; in females 32.3%. Hence, the influence of the host on the adult morphology of P. betularia, if present, is very small and re- mains below the resolving power of even this sophisticated technique. Zmales = Z females = Roskam: Platygaster and Metaclisis 129 Table 2. Summary of discriminant analyses. Values in brackets not used for calculation of canonical score Z. For further explanation, see text. CHARACTER canonical variates LARVAE length mandible diameter stigma Th 2 height tergal gland MALES width head length gaster length ult. flagellomere length are flagellomere width arn flagellomere FEMALES width head length gaster length ult. flagellomere th length 4 flagellomere width ach flagellomere 7 IMMATURE STAGES The larval phase of many Platygastridae is characterized by hypermetamorphosis: there are two distinct larval stages, which are very different in shape (Leiby & Hill, 1924). First instar larvae of forms in which hyper- metamorphosis occurs have a cyclopoid shape (figs. 21-23). They consist of a cephalothorax with huge mandibles to which a slender, 5—7 segmented abdomen is attached. Antennae are simple, conical. The surrounding of the mouth is sclerotized and differently shaped in the var- ious forms (figs. 18—20). Maxillary sensillae are only distinct in the form attributed to Piesto- pleura (fig. 22). In this form the cephalothorax bears two pseudopodia. In Metaclisis the ab- dominal segments are simple, whereas in cf. Piestopleura they seem to be secondarily subdi- vided. The final segment of Metaclisis is bilobed, coefficients for percentage of percentage of contribution variation load (0.04) (36.01) 84.43 27.46 5.44 in cf. Piestopleura it is simple, with its surface covered with small spinules. Stigmata are lack- ing. Platygaster does not pass a cyclopoid stage. In this genus, the final larval stage is preceded by a peculiar V-shaped structure (fig. 25). In the central “nodule” of this structure the embryo apparently develops, whereas the two arms of the “V” may function as teratocytes, structures immobilizing the endocrine system of the host and/or immunizing its encapsulating relations (Salt, 1968; Vinson & Iwantsch, 1980). The evi- dence, that these V-shaped structures do not be- long to the normal development of the host Se- mudobia is that in some instances the nodule becomes encapsulated by melanin. Although some extensive reports on early developmental stages of Platygaster exist (Marchal, 1906; Sil- vestri, 1916; Leiby & Hill, 1924; Hill & Emery, 130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 Figs. 18—24. Larval platygastrid characters. 18—20, detail of oral region; 21—23, cyclopoid larval stage, ven- tral aspect; 24, full grown larval stage, lateral aspect. 18—22, endoparasitoids of Dasineura interbracta; 23, Metaclisis phragmitis; 24, endoparasitoid of Semudobia brevipalpis. a, antenna; cl, clypeal sensilla; dp, dorsal protuberance; lbr, labral sensilla; m, mandible; mx, maxillary sensilla; prl, lateral prelabial sensilla; prm, median prelabial sensilla; ps, pseudopodium; s, stigma; sp, spine-like outgrowth; tg, tergal gland. 23, 24 x 100; 21, 22, x 240; 18—20, x 400. 1937; Clausen, 1956), V-shaped structures, in connexion to platygastrid parasitation, re- mained unobserved. After a moulting, final instar larvae develop from these primary stages. The terminology of the structures has been treated by Roskam (1982). The final larval stage is apodous and consists of a head, three thoracic segments (Th 1—3) and seven or eight abdominal segments (A 1—8) of which the final one is the anal segment (AS). The antennae are simple and inconspicu- ous. The clypeus bears one pair of papillae on which a seta may be developed. Mandibles are distinct. No sensillae are visible on the underlip complex. Second and third thoracic segments, and the second abdominal segment bear a pair of func- tional stigmata (figs. 26—27); the first abdomi- nal segment, and in Metaclisis also the abdomi- nal segments A 3—5, bear an oval, shallow, plate-like structure, in the centre of which a ves- tigial stigma is present (fig. 31). Silvestri (1916) made histological cross sections of these struc- tures and named them tergal glands. He sup- posed these glands to have a function during pu- pation of the parasitoid. Rows of papillae are present on dorsal, pleural and ventral surfaces of the body segments. Final instar larvae of Metaclisis, the two Platygaster species and the Nearctic form differ as follows. Metaclisis phragmitis (figs. 26, 30, 33, 35— 36). — Clypeal papillae with short seta. Mandi- bles small and curved (fig. 30). Eight abdominal segments present, with tergal glands on A 1, A 3—5. Papillary pattern rather complete, with one pair of rows of dorsal papillae, one pair of rows of pleural papillae and one pair of rows of sternal papillae (on thoracic segments) and ven- tral papillae (on abdominal segments). Two pairs of terminal papillae on dorsal surface of the anal segment. Dorsal papillae lacking on A 6 and A 7, pleural papillae sometimes doubled on Th 2 and Th 3. Ventral body surface with rounded verrucae. Platygaster (figs. 27, 28, 31, 32, 34). — Cly- peal papillae without seta. Mandibles straight and about twice as large as those of Metaclisis (fig. 28). Seven abdominal segments present, with tergal glands on A 1 only. Papillary pattern reduced and variable. Dorsal papillae usually absent. Pleural papillae only on Th 2 and Th 3 and, two pairs, on A 7. Sternal papillae, Roskam: Platygaster and Metaclisis 131 Figs. 25—36. Larval platygastrid characters. 25, third instar host larva with V-shaped endoparasitoid stage, lateral aspect; 26, 27, full grown endoparasitoid larva, latero-ventral aspect; 28—30, mandible of full grown en- doparasitoid larva; 31, tergal gland on first abdominal segment of full grown larva; 32, 33, stigma on second thoracic segment of full grown larva; 34, 35, head and sternal aspect of full grown larva; 36, ultimate and penul- timate segments of full grown larva, ventral aspect. 26, 30, 33, 35, 36, Metaclisis phragmitis; 25, 27, 28, 31, 32, 34, Platygaster betularia; 29, platygastrid endoparasitoid of Semudobia brevipalpis. a, antenna; cl, clypeal sensil- la; d, dorsal papilla; m, mandible; p, pleural papilla; s, sternal papilla; t, terminal papilla; tg, tergal gland; v, ven- tral papilla. 25, x 50; 26, 27, x 60; 34—36, x 150; 28—33, x 720. 132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 Table 3. Mean values (x) and standard deviations (s) of larval characters, measurements in um. Metaclisis, Platy- gaster and a Nearctic platygastrid. CHARACTER METACLISIS length mandible diameter stigma Th 2 height tergal gland sometimes doubled (fig. 34), on all thoracic seg- ments, ventrals only on A 1. Scattered spinules developed on dorsal surface of AS. Ventral sur- face of body segments with longitudinal striae. Nearctic form (figs. 24, 29). — No papillae visible, neither on clypeus, nor on body seg- ments. Mandibles straight and resembling those of Platygaster (fig. 29). Seven abdominal seg- ments present, with tergal glands on A 1 only. Anal segment with a huge, heavily sclerotized spine-like outgrowth (fig. 24: sp). In lateral as- pect the larvae are triangular, by a bizarre, allo- metric enlargement of the median protuberance between A 1 and A 2. Body surface without cu- ticular sculptures as verrucae and striae; ventral surface of A 5 slightly sclerotized. Means and standard deviations of the length of mandibles, the diameter of the stigma of Th 2 and the height of the tergal gland of A 1 are pre- sented for Metaclisis, Platygaster and the Nearc- tic form in table 3. Table 4 presents the same va- riates for larvae of Platygaster, dissected from Semudobia betulae (= P. betularia) and S. tarda (= P. betulae). Contrary to the results regard- ing intergeneric discrimination, only the height of the tergal gland provides discrimination at the species level. As in adults, discriminant anal- ysis provides the best separation between larvae PLATYGASTER NEARCTIC FORM of P. betulae and P. betularia, although some overlap (6.36%) remains. In table 2 a summary of values of the character set is given. The height of the tergal gland contributes most to the discrimination of the two species. In other words, specimens with high values for the, height of the tergal gland and low values for the length of the mandibles belong to P. betulae, whereas specimens with the inverse combina- tion of values belong to P. betularia. The ca- nonical score from the discriminant analysis is plotted in fig. 41. New specimens may be iden- tified by calculating their canoncial score Z as follows: Zlarvae = (height tergal gland) + 0.94 (diameter stigma Th 2) 0.64 (length mandible) > 80 P. betulae i esoyers = oem Spree P. betularia Because no discrimination was possible be- tween adult subgroups of P. betularia that de- veloped in the different host species S. betulae and S. skuhravae, such an analysis for the larval stages was omitted. PHENOLOGY, GEOGRAPHICAL DISTRIBUTION AND FURTHER BIOLOGICAL NOTES Phenological observations were made during 1972 and 1985 (figs. 42 and 43, respectively). Because the early larval stages of Semudobia Table 4. Mean values (x) and standard deviations (s) of larval characters, measurements in um. Platygaster spe- cies. (nd), overlap very large, not defined. CHARACTER betularta length mandible diameter stigma Th 2 height tergal gland betulae % overlap Roskam: Platygaster and Metaclisis 133 Figs. 37—40. Adult scutellum, propodeum and first gastral tergite. 37, Platygaster betularia, male; 39, ditto, female; 38, P. betulae, male; 40, ditto, female. x 250. species and Platygaster species are difficult to identify during mass inspections, the 1972 re- sults are not presented for the separate species. The 1985 results, specified for the species in- volved, show that the interspecific differences concerning the flight period are small, for Se- mudobia, as well as for Platygaster. P. betula- ria, the most abundant parasitoid in Meijendel, appeared first, followed by P. betulae. M. ph- ragmitis is the last one, but differed only five days with P. betularia. All parasitoid species have a considerably longer flight period than their hosts. The slight difference between P. be- tularia and P. betulae is corresponding to the difference of maximum activity of their re- spective hosts, S. betulae and S. tarda. Adult stages of gall midges, as well as of para- sitoids, appeared about a fortnight earlier in 1972 than in 1985, probably due to the very cold spring of the latter year. Furthermore, adult gall midge activity lasted considerably longer in 1972 than in 1985, as did, to a lesser extent, the activity of the platygastrids. A possi- ble explanation for the latter difference may be the great variation of the maximum temperature in 1985: a short period of very warm weather (17—19 May; tmax = 25°C) was followed by an extraordinarily (23—24 May; tre DO). Platygaster — In the field, adult emergence coincides with the appearance of Semudobia fe- males. Ovipositing gall midges and parasitoids frequently occur together on the same flowering birch catkin, but in other instances Platygaster max cold period 134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 20 Q | © . | LARVAE x | 15 O | e e | ® 10 e | | * * | * | * 5 * > * Kk | * A (o) | | FSO TE ER WR eh SIONI VUE SEE Aigo OL “5 “4 -3 2 “a 0 1 2 SCA 5 6 | DA 1 20 e | © | e © MALES je 5 | . È | e e 10 * | à | * * | * 5 * | ni | * * | * o | ET SR I aN RA "6.25 5.00 375 “2.50 -1.25 0 125 250 375 5.00 6.25 7.50 | DA 1 20 e | bd À © FEMALES ® 15 e © e ® e 10 * * * * | * 5 * | = | * * | * al | | "6-25 “5-00 “3-75 "2-50 1-25 0 1-25 250 375 500 6-25 7-50 DA 1 Fig. 41. Two group discriminant space. Y-axis represents only the sequence of specimens. DA 1, first discrimi- nant axis; dots, Platygaster betulae; asterisks, P. betularia. Roskam: Platygaster and Metaclisis 9 16 23 30 6 13 20 27 4 11 18 25 1 8 TR cee UU Be ae al TI EED | 5 CET Pupa GED adult GED ©9099 198227729756 135 13 20 27 3 10 17 24 31 7 14 21 ar VU ear U dert Visita (Vite rl Sie. 2 Bon ha ERRATA] CEE REG SPEER ETSEN | CRASS |; SEMUDOBIA | ||| pupa GED adult PLATYGASTER || pupa GED adult METACLISIS ee Lo) TEEN FRE) pupa @ \V-shaped EEE ER REED Lu (GE pupa «> cyclopoid EB L, Fig. 42. Phenology of platygastrıd parasitoids, Meijendel, 1972. L1, L2, L3, Lm: first, second, third, full grown larval instar, respectively. females search for host eggs in absence of Semu- dobia. Adult parasitoid activity ceases when most of the host eggs have been eclosed in the last week of May. Parasitized gall midge em- bryo’s apparently develop in a normal way. Hatched host larvae with dormant parasitoids mine into ovaries of Betula and induce galls as do healthy larvae. Not before the host reaches its early third instar, signs of parasitation ap- pear. The host becomes less mobile and looses its bright orange colour. In this stage the V- shape “teratocyte stage” becomes apparent. From the end of June until mid-August the final instar larva fills about the whole body content of its host. From the end of July parasitoids pu- pate, remaining within the host skin and filling about half the room with meconium. In the sec- ond half of August the pupa is dark, fully scle- atized and looses its, exuviae ‘The adult over- winters in a quiescent condition and leaves the gall when the temperture rises at the end of April of the following year. Metaclisis. — This parasitoid is about a week later in development than Platygaster. The peri- od of adult flight is somewhat shorter than that of both Platygaster species; it lasts only two to three weeks. Cyclopoid larvae of Metaclisis be- come visible when the host is in its early third (final) instar, as does the V-shaped stage of Platygaster. The cyclopoid stage, however, lasts considerably longer than the V-shaped one. Platygaster and Metaclisis were present in all samples collected in North-Western Europe, Switzerland and Poland. Platygaster was also reared from samples collected in Wladiwostok, U.S.S.R. and Sapporo, Japan. The lack of Meta- clisis in these samples may be attributed to small sample sizes. Parasitoids belonging to Platygas- ter and Metaclisis are absent from the Nearctic: over 70 samples collected in Canada (Alberta and Quebec) and U.S.A. (Pennsylvania, Ohio, Illinois, South Dakota, Montana, Wyoming and Colorado) contained abundant Semudobia galls, but were free from these parasitoids. The un- described platygastrid endoparasitoid was dis- sected from galls induced by S. brevipalpis Ros- kam in fruit catkins of Betula populifolia Marsh. (Pennsylvania, Catskill formation, Long Pond, 136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 P. betulae Q Fig. 43. Field captures of adult platygastrid parasitoids, Meijendel, 1985. Drawn lines, parasitoid females; dot- ted lines, host females. Luzerne County, Leg. A. A. Heller & E. Ger- trude Halbach, 16—17.1x.1892). Not only egg- larval parasitoids, one sample with the undes- cribed form excepted, but also inquiline gall midges are absent from Nearctic samples (Ros- kam, 1979). Hence, Nearctic insect commu- nities centered upon Semudobia are less diverse than Palaearctic ones: two complete segments of the food web, including important mortality factors in the Palaearctic, are missing in the Nearctic. Endoparasitoids of Dasineura. — Two gall midge species, viz., D. fastidiosa Roskam and D. interbracta Roskam occur frequently in birch catkins. They are inquilines (food parasi- toids) of Semudobia species (Roskam, 1979). Cyclopoid stages of platygastrid endoparasi- toids (figs. 21, 22) were dissected from Dasineu- ra larvae collected from 7.vi—3.vii.1978, Mei- jendel, and on 20.vii.1977, Norway, Aseral. Be- cause Dasineura larva drop onto the ground before the parasitoids reach the full grown instar, this stage has not been observed and adults could not be reared. In each parasitized Dasineura larva usually two parasitoids, attributed to Piestopleura, are present. The pairs of parasitoids are supposed to be twins as the result of polyembryonic devel- opment. A twinning development was earlier reported for another platygastrid, namely, Platygaster hiemalis Forster (Leiby & Hill, 1923). PARASITOID SPECIFICITY In order to determine host — parasitoid spe- cificity, adult parasitoids have been reared from sorted samples. The results are presented in ta- ble 5. Metaclisis phragmitis, although less common than both Platygaster species, is a regular para- sitoid of S. betulae and S. tarda, but could only be reared once from S. skuhravae. Platygaster Roskam: Platygaster and Metaclisis 137 betularia is a common parasitoid of S. skuhra- secting three samples of galls, collected in De- vae and S. betulae, whereas P. betulae is only cember, 1975 and 1983. The results are pre- abundant on S. tarda. Because egg sizes of the sented in table 6. | various host species are different (Roskam, Asa rule, mortality caused by Metaclisis phrag- 1977), parasitoids might be able to discriminate mitis remains low; only in 1983, Meijendel, between host eggs. Furthermore, host prefer- mortality of Semudobia betulae approached ence might also have a phenological basis, be- 10%. Platygaster, however, contributed consid- cause small, but consistent differences exist be- erably to gall midge mortality. In Duiven- tween the phenologies of the host species (Ros- voorde, 1975, and Meijendel, 1983, P. betulae kam, 1977): S.skubravae emerges first, alone caused a higher mortality than all chalci- followed by S. betulae; S. tarda is usually the doid parasitoids (about four species, belonging latest. Metaclisis does not emerge before the sec- to three genera) together. ond week of May; eggs of S. skuhravae may not Densities (numbers of specimens per unit of be appropriate then anymore for oviposition of area) of hosts and parasitoids may be interre- this parasitoid. For the same reason the pheno- lated. Whether density-dependent effects exist logies of P. betularia and S. tarda may not and how to determine these effects has been match and, on the other hand, those of P. betu- treated by Southwood (1978). Many reports on lae, S. skuhravae and S. betulae. this subject have been discussed by Stubbs Within the large genus Platygaster, P. betulae (1977). Southwood (1978, and references and P. betularia belong to different species therein) supposed an exponential interdepen- groups, which Kieffer (1926) considered as dif- dence between the original density of host pop- ferent genera. This implies that the closest rela- ulation N, and the density of survivors N, of a tives of both species did develop on other hosts particular mortality factor according the func- than Semudobia. Therefore, the ecological asso- tion. ciation between Semudobia and Platygaster did N, = A(N,)8 (1) not affect their respective speciation patterns where A and B are constants that define the and a co-evolutionary process cannot be re- relationships between mortality and density. In sponsible for host- and egg-larval parasitoid di- logarithmic form the equation is linear versity. This is probably in contrast with the Log N, = log A + Blog N, (2) speciation patterns of the food parasitoids, viz., where B defines the slope of the regression line Dasineura interbracta and D. fastidiosa (Ros- of log N, over N,. When B does not depart sig- kam, 1979), and some of the chalcidoid parasi- nificantly from 1, a density-dependent effect is toids (Roskam, in preparation). absent. However, when B < 1, the mortality factor has a positive density-dependent effect: . high host densities (aggregated situations) suffer Host mortality has been determined by dis- proportionally more than low densities. B > 1 HosT MORTALITY Table 5. Parasitoid specificity regarding various Semudobia hosts. a na a a a a SEMUDOBIA SKUHRAVAE 9 SEMUDOBIA BETULAE ° SEMUDOBIA TARDA ° E E = ae 2 SE 3 ÿ © x à È $ ue Da 8 È < = 8 = = x 3 3 = 3 » » a IS] » » a 8 » a ae. UE NE STERN a SES 8 à à À 8 à 2 È SUR (©) o) . È (©) . oO . . Le] = A A, = = 2 = Ay 2 x "n x Le] oO oO Hulshorst Kootwijk 21 Meijendel 57 213 15 46 303 Meijendel Duivenvoorde 76 4 Sf om = 8 29 73) 138 14 Ilperveld 1979 Nieuwkoop 38 SENTE ie A erie: I, 207502 [aus 1 - 26 116 1 sl walt = 86 | 388 6 ils} 5 - 110} 134 34—43 - - 90—293 1073/1827 49—47 92—183 1—9 roses 138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 Table 6. Mortality (%) caused by parasitoids. —, not defined. skuhravae Duivenvoorde Meijendel Meijendel betulae Duivenvoorde Meijendel Meijendel tarda Duivenvoorde Meijendel Meijendel represents the inverse situation: density depen- dence is negative, low host densities (segregated situations) become proportionally more se- verely attacked. The intercept of the regression line, log A, is not further considered; the mean- ing of this constant is discussed by Hassell (1975). In order to determine the density-dependent of GALLS PLATYGASTER METACLISIS CHALCIDOIDS INQUILINES NR. eftect of Platygaster, two samples of catkins (ten per tree) were collected in Meijendel, December 1982 and 1983. Because each catkin was consid- ered a functional unit of area, a patch, the num- bers of galls (N,) and of galls without Platygas- ter parasitation (N,) were defined per catkin. Interference between Platygaster and other par- asitoids was not considered because chalcidoid Table 7. Density-dependent host mortality. (*), significant, p < 0.05. GALLS per CATKIN skuhravae 1982 1983 betulae 1982 1983 1982 1983 of CATKINS NR. with GALLS of CATKINS with PLATYGASTER of GALLS in EMPTY PATCHES NR. of GALLS in LOCALIZED PATCHES NR. NR. Roskam: Platygaster and Metaclisis 189 parasitoids either refuse host larvae with Platy- gaster parasitoids (adults), or perish on such hosts (larvae). Galls attacked by inquiline Dasi- neura, which indeed may contain parasitized Semudobia larvae, have not been considered too, because it appeared impossible to deter- mine parasitation of such hosts. Two aspects become distinct from an analysis of the results (table 7). First, gall midge densities vary considerably among different years. S. skuhravae, as well as S. tarda, caused in 1983 a tenfold of the galls of the preceding year and S. betulae produced four times more. Large dif- ferences among generations of different years have also been found for other gall midge spe- cies and may occur commonly (Skuhrava et al., 1984). The mechanisms that cause such large fluctuations are not well understood. Second, Platygaster species indeed have dif- ferent effects upon different gall midge densi- ties. Only about one third, or less, of catkins with galls in low densities (those of S. skuhra- vae and S. betulae, 1982), but about two third of catkins with galls in high densities (S. tarda, 1983) contained parasitized gall midge larvae. This means that many low density patches re- mained unnoticed (,,not localized”) by oviposit- ing parasitoids and may therefore function as escape possibilities for the midges. In high den- sity situations (S. tarda, 1983), almost all galls occur in catkins found by parasitoids. If B-val- ues are defined for patches, visited by parasi- toids, a rather surprising result emerges: either B does not significantly depart from 1 (S. betu- 4 2.00 1.75 Log N, o Log N, = 7 0.2547 +1.1338 log N, 0.2 0-6 1.0 1.4 1-8 2-2 lae — P. betularia, 1983; S. tarda—P. betulae, 1982), which means that the parasitoids are un- able to regulate the host densities; or B is signif- icantly larger than 1 (S. betulae—P. betularia, 1982; S. tarda—P. betulae, 1983; fig. 44), which means that in those cases the parasitoids have a negatively density-dependent impact on their hosts. Moderate (and, when localized, low) host densities suffer more than high ones. Hence, in localized catkins, escape possibilities for the midges are larger in patches with high densities. Combining the outcome for localized and not localized catkins, the conclusion is that Platy- gaster parasitoids may be able to eliminate mod- erate host densities. Escape possibilities for the gall midges remain in both tails of their density distribution: in highly segregate, as well as in highly aggregate situations. CONCLUSIONS 1. Adult Platygaster betularıa and P. betulae can be distinguished by a combination of anten- nal characters; larvae by a combination of char- acters regarding the height of the tergal gland, “the diameter of the stigma on the second thorac- ic segment and the length of the mandibles. 2. No discrimination is possible between subgroups of a parasitoid species that developed in different host species (e.g. P. betularia reared from S. skuhravae or S. betulae). 3. All platygastrid egg-larval parasitoids de- velop highly synchronized with their hosts. Metaclisis phragmitis develops about one week later than both Platygaster species. This pheno- Ya Log N, = 0.3117 + 1.1228 log N, 0-2 0-6 1.0 1.4 1-8 2.2 Log N; Fig. 44. Density-dependent host mortality. B, slope of the regression line; N,, density of survivors; N,, original density of host population. Drawn line, Du cn of N, over N,; dashed fae = dele graph, Platygaster betularia on Semudobia betulae, Meijendel, 1982 ther explanation, see text. ; right graph, Je. penne on S. tarda, Meijendel, 1983. For fur- 140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 4, 1986 logical difference might explain the absence of Metaclisis phragmitis from S. skuhravae. 4. Platygaster has been reared from Western and Eastern Palaeartctic localities. Platygaster and Metaclisis are absent from Nearctic Semu- dobia galls. One Nearctic collection of an un- known egg-larval parasitoid excepted, the whole guild of egg-larval parasitoids is absent from this region, as is the guild of inquilines. 5. Almost complete separation exists in the host preference of the two Platygaster species: P. betularia is a common parasitoid of S. skuh- ravae and S. betulae, whereas P. betulae is com- mon on S. tarda. Metaclisis phragmitis, one col- lection excepted, has not been reared from S. skuhravae. 6. Diversity of egg-larval parasitoids and their hosts is not a result of co-evolution, be- cause P. betularia and P. betulae belong to dif- ferent species groups, whereas Semudobia spe- cies are close relatives. 7. Platygaster species are important mortali- ty factors of Semudobia species and may have a density-dependent impact on their hosts. Es- cape possibilities for Semudobia are largest in highly aggregate situations, as well as in highly segregate ones. ACKNOWLEDGEMENTS I thank Mr. H. J. Vlug, Wageningen, for con- firming the identifications and for critical read- ing of a draft of this paper. Dr. M. Zandee, Lei- den, provided the statistical advice and Mr. K. Jalink prepared the SEM photographs. REFERENCES Bachmaier, F., 1965. Die Insekten- und Milbenfauna der Zwergbirke. — Veröff. zool. St. Samml., Münch. 9: 55—158. Barnes, H. F., 1951. Gall midges of economic impor- tance. V. Trees: 270 pp. — London. Bigelow, R. S., & C. Reimer, 1954. An application of the linear discriminant function to insect taxono- thy. — Can. Ent. 86: 69—73. Clausen, C. P., 1956. The egg-larval host relationship among the parasitic Hymenoptera. — Boll. Lab. Zool. Gen. Agr. Portici 23: 119—133. Debauche, H. R., 1947. Scelionidae de la faune belge — Bull. Annls. Soc. r. ent. Belg. 83: 255—285. Fulmek, L., 1968. Parasitinsekten der Insektengallen Europas. — Beitr. Ent. 18: 719—952. Hassell, M. P., 1975. Density-dependence in single- species populations. — J. Anim. Ecol. 44: 283— 295. Hill, C. C., & W. T. Emery, 1937. The biology of Platygaster herrickii, a parasite of the hessian fly. —]. a Res. 55: 199—213. Hodges, S., 1969. Gall midges (Diptera-Cecidomyii- Piremal KR. A., dae) and their parasites (Hymenoptera) living in female birch catkins. — Trans. Soc. Br. Ent. 18: 195—266. Kieffer, J. J., 1916. Beitrag zur Kenntnis der Platygas- terinae und ihrer Lebenweise. — Centralbl. f. Bakt. 46: 547—592. Kieffer, J. J., 1926. Scelionidae. — Das Tierreich 48: 1—885. Leiby, R. W., & C. C. Hill, 1923. The twinning and monembryonic development of Platygaster hie- malis, a parasite of the hessian fly. — J. Agric. Res. 25: 337—350. Leiby, R. W., & C. C. Hill, 1924. The polyembryonic development of Platygaster vernalis. — J. Agric. Res. 28: 829—840. Lubischew, A. A., 1962. On the use of discriminant functions 1 in taxonomy. — Biometrics 18: 455— 476. Marchal, P., 1906. Recherches sur la biologie et le développement des Hymenopteres Parasites. II. Les Platygasters. — Arch. Zool. Exp. Gén. (4° Ser.) 4: 485640. 1979. Morphometrics: 276 pp. — Dubuque, Iowa. Roskam, J. C., 1977. Biosystematics of insects living in female birch catkins. I. Gall midges of the genus Semudobia Kieffer (Diptera, Cecidomyiidae). — Tijdschr. Ent. 120: 153—197. Roskam, J. C., 1979. Biosystematics of insects living in female birch catkins. II. Inquiline and preda- ceous gall midges belonging to various genera. — Neth. J. Zool. 29: 283—351. Roskam, J. C., 1982. Larval characters of some eury- tomid species (Hymenoptera, Chalcidoidea). — Proc. Kon. Ned. Ak. Wet. (Ser. C) 85: 293—305. Roskam, J. C., & G. A. van Uffelen, 1981. Biosyste- matics of insects living in female birch catkins. III. Plant-insect relation between white birches, Betu- la L., Section Excelsae (Koch) and gall midges of the genus Semudobia Kieffer (Diptera, Cecido- myiidae). — Neth. J. Zool. 31: 533—553. Salt, G., 1968. The resistence of insect parasitoids to the defence reactions of their hosts. — Biol. Rev. 43: 200—232. Silvestri, F., 1916. Contribuzioni alla conoscenza bio- logica degli Imenotteri Parassiti. V. Sviluppo del Platygaster dryomyiae Silv. (Fam. Proctotrupi- dae). — Boll. Lab. Zool. Gen. Agr. Portici 11: 299—326. Skuhrava, M., V. Skuhravy, & J. W. Brewer, 1984. The distribution and long-term changes in popula- tion dynamics of gall midges on cereals in Europe (Cecidomyiidae, Diptera). — Cecid. Internation- ale 5: 1—5. Southwood, T. R. E., pp. — London. Stubbs, M., 1977. Density dependence in the life-cy- cles of animals and its importance to K- & r-strat- egies. — J. Anim. Ecol. 46: 677—688. Vinson, S. B., & G. F. Iwantsch, 1980. Host regula- tion by insect parasitoids. — Quart. Rev. Biol. 55: 143—165. 1978. Ecological methods: 524 DEEL 129 AFLEVERING 5 1986 IL 161 "568 = TIJDSCHRIFT UITGEGEVEN DOOR \ KD i WS, \ Sa = Sceliphron deforme tibiale Cameron (figs. 40, 116) Sceliphron formosum; Bingham, 1897: 236, 239, 2 d. Sceliphron tibiale Cameron, 1899: 53 (sex not men- tioned) — Khasia Hills (OUM; examined). Sceliphron lineatipes Cameron, 1900: 36, 2 & — Kha- sia Hills, coll. Rothney (OUM; examined). Sceliphron deforme; Strand, 1914: 116 (nest); Kohl, 1918: 122 (Bombay, Darjeeling, Burma: Moul- main, Tenasserim). R. V. Hensen: The subgenus Prosceliphron 235 55 57 Figs. 49—57. Sceliphron fervens (Smith). 49—50, 2, Indonesia, Borneo, 49, clypeus; 50, antennal insertion. 51, 2, Malaysia, propodeal orifice. 52—55, 3, Malaysia, 52, clypeus; 53, genitalia, ventral aspect; 54, inner side of left half of aedeagus; 55, metasoma, lateral aspect. 56, 2, Malaysia, pronotum, lateral aspect. 57, 2, Indonesia, Borneo, first metasomal segment. 49, 52, 55, 57: scale-line; 56: 2.0 X scale-line; 50—51: 4.0 X scale-line; 53, 54: 8.0 x scale-line. 236 Sceliphron (Prosceliphron) deforme tibiale; Bohart & Menke, 1976: 106. Types. — The OUM possesses one female specimen labelled “Sceliphron tibiale Cameron, type, Khasia”, which I herewith designate as the lectotype. In the BMNH I have seen two fe- males labelled “S. tibiale Cameron, Khasia”, “ex FMS 1955—354”, which I have labelled as para- lectotypes, since S. tibiale was described from material in the FMS-collection. S. lineatipes was described from OUM- material. I found there two male specimens on one pin, with label “Sceliphron lineatipes, type, Cameron”. The upper specimen fits the rather incoherent description best: it agrees at least with the Latin diagnosis. I herewith designate it as the lectotype. The lower specimen is paralec- totype, as is a male labelled “Sceliphron linea- tipes Cam., type, Khasia” in the BMNH. Sever- al of 72 partly unlabelled specimens in the OUM and ZMA may have been part of the or- gininal type-material too, since S. tibiale was described from both sexes, but there is no evi- dence based on label data which allows desig- nation of more paralectotypes. Description. — Post-antennal tubercles small (cf. fig. 50), black. Yellow colour-pattern rather extensive (fig. 40). Characteristic are: antennal scape yellow or reddish yellow; propodeum of- ten with a pair of spots at apex of the dorsal en- closure; legs reddish except the coxae, trochant- ers and base of femora dorsally; metasoma with broad yellow bands on all tergites, those on first and second tergite suffused with red or reduced; sternites with lateral reddish-yellow spots. In the male the coloration is like the female or more or less reduced: clypeus with a pair of small spots or completely black, scapes dorsally black, spots at basis of propodeum small or ab- sent; legs brownish, hind femora ventrally somewhat lighter; metasoma with narrow bands on fourth to sixth tergite, those on second and third reduced or absent. Variation. — Specimens from Darjeeling tend to be more brightly coloured than described above: in the male the clypeus bears a large mark, the scapes are red dorsally, the legs are mainly red-yellow and the metasoma has com- plete bands on all tergites. Material examined. — India. — 1 ®, Dacca, 30 May 1945, D. Leston (BMNH); 2 2, N. Khasia Hills (BMNH); 5 © 5 6, Khasia, coll. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Bingham (OUM); 4 @, Sikkim, Runjit Valley, 1000 ft., April 1894, Bingham coll. (BMNH); 3 2, Sikkim, F. A. Müller (UZM); 1 ©, N.W. In- dia (BMNH); 1 ©, N. India, Lukna, 55 km S. Darjeeling, May 1966, Sedlacek (BPBM); 3 ©, Sikkim, coll. Bingham (BMNH); 2 2 1 6, Dar- jeeling, coll. Frühstorfer (RMNH); 19 ® 10 8, Khasia Cameron (ZMA); 10 ©, Sikkim, coll. Bingham (ZMB). Burma. — 1 2, Rangoon Dist., March 1888, Bingham coll. (BMNH); 1 2, Pegu Hills, March 1889 (BMNH). Sceliphron deforme femorale ssp. n. (figs. 38, 41, 116) Holotype. — 2, Thailand, Chiangmai, 4 May 1952, D. & E. Thurman (USNM). Description. — Post-antennal tubercles small, black (like in S. d. tibiale); colour-pattern much less extensive than in adjacent subspecies (fig. 41); antennal scape yellow ventrally; pronotum with narrow, interrupted yellow band; subtegu- lar spots small; scutellum with narrow trans- verse spot; propodeum only with small apical spot; fore and middle femora black, with small yellow spot apically; hind femora reddish; legs otherwise blackish; metasoma black, first tergite medially reddish-yellow, third with strongly re- duced, fourth with complete yellow apical band; fifth and sixth reddish-yellow; tomentum of face golden, erect pubescence purely white. Structurally S. d. femorale is identical with S. d. tibiale. With respect to colour-pattern how- ever, the form resembles the related species S. fervens, which occurs in the adjoining area; this particular colour-pattern may thus represent an independently developed environmental adapta- tion. Paratypes. — Thailand. — 1 ©, Doi Suthep, 4—10 May 1952, D. & E. Thurman (USNM); 2 2, Khun Tan Mts., May 1933, 4000 ft, H. Smith coll. (USNM, RMNH) (with basal spots on propodeum, larger scutellar band, one with complete band on third tergite). Laos. — 2 9, Pak Kop, Mekong River, 7 May 1920, R. V. de Salvaza (BMNH) (one of these with propodeal spots basally). Sceliphron deforme deforme (Smith) (figs. 31, 43—45) Pelopoeus deformis Smith, 1856: 231, ® — North China, coll. Fortune (BMNH; examined). Sceliphron deforme; Strand, 1914: 116 (Tsingtau, 237 R. V. HENSEN: The subgenus Prosceliphron -uemegeg ‘sourddryryg “è ‘09 “years ‘eiskefew “6G SPISAETETN 152 M ‘gg 'suraned 1n0]09 “(rung ) suaaraf U014d1722S ‘09—8G “SSI 238 nest); Turner, 1917: 176; Kohl, 1918: 122—123, figs. 26, 27; Gussakovskij, 1936: 4 (N. E. Szech- wan); Iwata, 1939: 169, fig. 21 (Formosa, bionom- ics); Yasumatsu, 1942: 106 (Peking); Tsuneki, 1967 (Formosa). Sceliphron deforme taiwanum Tsuneki, 1971: 6, 2 d — Taiwan, Taitung pref., Chulu (coll. Tsuneki; not examined); Bohart & Menke, 1976: 106. Sceliphron (Prosceliphron deforme deforme; Bohart & Menke, 1976: 106. Type. — The specimen which was figuring as the type of S. deforme in the BM (no. 21.613) belongs to S. curvatum. It was evidently incor- rectly labelled, since it does not agree with the description of S. deforme, which mentions a “spot at each side of the metathorax at base”; such spots fail in S. curvatum, and moreover, S. curvatum does not occur in Northern China. The lectotype by present designation is a female in the BMNH with label “N. China, 54—8” (according to the register: “China, Shanghai, purchased from Stevens, coll. by Fortune”). There is one other specimen which is old enough, and may have been a syntype. It is la- belled “56—45, China” (“April; China, purch. from Stevens, coll. by Mr. Bowring”) and “type rufopictus Smith” in Smith’s handwriting. There are two female specimens in the Saun- ders’ collection (OUM) standing as syntypes of S. deforme. One of these belongs to S. deforme, the other to S. curvatum. There is no evidence that these were part of the original type-materi- al. Description. — Female. — Post-antennal tu- bercles of intermediate size (fig. 31), larger than in S. d. tibiale, smaller than in S. d. atrıpes, usually with yellow spot; colour-pattern rather extensive (figs. 42—44), only slightly darker than tibiale: scapes often black dorsally, tegulae with yellow mark, propodeum with large basal spots, and sometimes with a pair of spots at apex of the dorsal enclosure, legs blackish to red-brown, femora largely black dorsally, ven- trally lighter, metasoma with broad reddish-yel- low bands on alle tergites, sternites 2—5 with bands or lateral spots. Male. — Darker than the female, but very variable: clypeus often only with a pair of small spots, basal propodeal spots reduced or absent, bands on tergites usually reduced, sometimes nearly absent. Variation. — Female specimens from Tibet are very brightly coloured (fig. 42): the scapes are completely yellow, post-antennal tubercles TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) with yellow mark, often small spots on sides of | pronotum, and marks on the mesoscutum in front of the tegula, propodeum with a pair of spots in the dorsal enclosure, legs mainly yel- low, femora dorsally black on proximal half, pe- tiolus largely or partly yellow. The coloration of the Taiwanese population was described by Tsuneki (1971), and is illustrated in fig. 44. The pattern is somewhat less bright than that of the Tibetan specimens, and brighter than that of the other mainland populations (fig. 45). I prefer therefore to regard Tsuneki’s S. deforme tai- wanum as part of the nominate species. Material examined. — Mongolia. — Tchi-Li, J. de Joannis, 1890 (MNHN); Tcheli, J. de Joannis, 1903 (MNHN). China. — 1 ?, Ningpo, Aug. 1949 (NMB); 1 2, China (OUM); 2 2, Tientsin, N. China; 1 ES Le > ©, Peking, “curvatum, det. Kohl”, “deforme, . det. Kohl”; 1 ?, Tientsin, Walte; 1 9, Tsingtau, Prof. Hoffmann; 1 2, Tschili; 1 6, Tientsin (all NMW); 3 & 1 6, Shankhov, Honan, D. Renard (IRSN); 4 © 2 g, Kiangsi, A. David, 1875; 1 9, Kiangsu, Shanghai, 27 July 1925, 1 d, Shanghai, 14 Aug. 1924, both O. Piel; 1 d, Kouytcheou, 1921, Cavalérie; 1 2 1 d, Shensi, 1875, A. Da- vid (all MNHN); Canton, 1 ©, 8 June 1917, 28 1 2, 10 May 1917; Lungtaoshan, Kwangtung, 1 2, 17. Sep 19171722719 Sep. MRC PIRE 1917; Lofaoshan, Kwangtung, 1 ®, 20 July 1916, 1 d, 11 Aug. 1916, 1 2 1 d, 20 May 1917, 1 4,1 May 1917; Gaotung, Kwangtung, 1 d, 3 June, 1 6, 9 June; Tshayunshan, Kwangtung, 1 2, iil Aug. 1912, 1 2,24 June IDEE June 1912; 1 ?, Sholyunshan, 30 May; all leg. Mell (ZMB); 1 2, Chinkiang, 10 June 1924, 1 2, Mokanshan, 18 July 1924, both J. F. Illing- worth (BMNH); 1 ©, Hongkong, F. W. Ferry, 1 2, Tonkin, Hoabink, Aug. 1918, R. V. de Sal- vage; 2 ©, Howlik, R. C. L. Perkins coll.; 1 ®, Peiping, C. F Wu coll.; 1 2, Foochow, June 1936, M. S. Yang; 1 2, Yunnan, 1918, G. For- rest; 1 2, Shanghai, 1 ©, Sinling, pr, Shen-Se, W. A. Maw; 1 6, Shihchiachwang, C. F. Wu coll. (all BMNH); 5 @, Kiangsu, Kolthoff (NRS); 1 ©, Kina, N.O. Szechwan, Sven He- dins Exp. C. Asien, Dr. Hummel (NRS); 1 2, Foochow, Kellogg (MCZ); 1 ©, Huaying Shan, Szechwan, Aug. 1932, G. Liu (MCZ); China- Tibet border, 3 2, Ludingshiao, 4900 ft., 12— 20 Aug. 1930, 1 2, Bet. Uenchuan and Mow- chow, 4500—5500 ft., 3—6 Aug. 1924, 1 = Dashianglin Pass, 4960 ft., 23 Aug. 1930; Suifu, 178722,1928, 1 27 May-June 192951008 R. V. HENSEN: The subgenus Prosceliphron 239 65 66 68 1.0 mm 67 Figs. 61—69. Sceliphron coromandelicum (Lepeletier). 61—62, 6, Malaysia. 61, genitalia, ventral aspect; 62, in- ner side of left half of aedeagus. 63—65, ©, India, Kerala. 63, antennal insertion; 64, propodeal orifice: 65, cly- peus. 66—67, 3, Malaysia. 66, clypeus; 67, first metasomal segment. 68—69; 2, Malaysia. 68, first metasomal segment; 69, pronotum, lateral aspect. 65—68: scale-line; 69: 2.0 X scale-line; 63, 64: 4.0 X scale-line; 61, 62: 8.0 x scale-line. 240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) 2000 ft.; 5 2, Shinkaisi, Mt. Omei, 4000—5000 ft) ee Meo Er LOO vir som lyn 1929 MRC Chiacopin, 22 Jan. 1933, 2100 ft., 1 2, Dono- menwai, 10 mi. W. Weichow, 5600 ft., 1933; 1 2, S. of Suifu, Aug. 1929; 1 ©, Tseojiageo, S. of Suifu, Sep. 1929, 1400—2000 ft.; 1 9, near Ya- chow, 2 July 1930; 7 ®, Kuanshien, 3000 ft., 1-4 Aug. 1934; 8 22 6, Fulin, 2400 ft., 17— 18 July 1928, 2 2, Ningyuenfu, 6000—6200 ft., 2-4 Aug. 1928; 2 ©, Bet. Ginkeoho and Shin- kaishi, 3—4 Aug. 1925; 1 ©, Tsaekeo; all D. C. Graham coll. (USNM). Taiwan. — 1 ©, Taihorinsho, Aug. 1908, 1 ®, Fuhosho, Sep. 1909, 2 ®, Taihorinsho, Oct. 1909, 2 ©, Fuhosho, Aug. 1909, 1 ©, Kanshirei, 108 a AS ate RNA MS rar AZ [ay 1934, L. Gressit (MCZ); 6 d 1 ®, Taipei-hsien, Wulai, 4 July 1966, 1 4, Nantou-hsien, Puli, 11 July 1966, 3 2, Taitung-hsien, Chulu, 12 Aug. 166, I 2, ick, 29 one 1968, 2 2, Taitung- hsien, Chihpechi, resp. 13 Aug. 1966 and 30 June 1968, all leg. K. Tsuneki (RMNH). Japan. — Chiba Pref., 1 ?, Usui, Sakura City, 9 Sep. 1968, 1 2, Oamish irasato, Sanbu Gun, 10 Sep. 1970, 1 ®, Kuriyama, Matsudo City, 22 Aug. 1963, 2 d, Mt. Kasamori, Cho- nan, 3 Sep. 1970, 1 d, Zenshoji Toke, Chiba City, 10 Sep. 1970, all leg. H. Suda (RMNH). Vietnam. — 2 ®, Ht. Tonkin, Phuong Lom, (J. J. M. Laisi) R. Oberthir 1901 (MNHN) (transitional between S. d. deforme and S. d. femorale; band on second tergite reduced, hind femur less dark than fore and middle). Sceliphron deforme atripes (Morawitz) (figs. 32, 46, 116) Pelopoeus atripes Morawitz, 1888: 271, 2 — Semipa- latinsk (ZIL; examined). Sceliphron deforme; Kohl, 1918: 122, 123, 133 (origi- nal description). Sceliphron formosum var. koreanum Uchida, 1925: 329 (description in Japanese) (not examined). Sceliphron deforme koreanum; Tsuneki, 1967: 6 (de- scription 9 ó). Sceliphron (Prosceliphron) deforme atrıpes; Bohart & Menke, 1976: 106. Sceliphron (Prosceliphron) deforme koreanum; Bohart & Menke, 1976: 106. Type. — The Zoological Institute at Lenin- grad possesses two syntypes of P. atripes Mora- witz. The lectotype, by present designation, is a female, labelled “Semipalatinsk”, “k. F. Mora- witz” and “atripes 2, F. Morawitz”. The para- lectotype ıs labelled “Semipalatinsk”, “k. F. Morawitz” and “Pelopoeus atripes ©, F. Mora- witz”. It lacks forewings, antennae and the legs largely. Description. — Morphology. — Post-anten- nal tubercles exceptionally large, dorsally trun- cate (fig. 32). Coloration. — In the type-specimens, the colour-pattern is strongly reduced (fig. 46): the metasoma bears no yellow bands, and the meso- soma is almost entirely black; specimens from the Eastern part of the area occupied by this subspecies are more brightly coloured (fig. 47). Material examined. — Mongolia. — 1 2 1 6, Dinyuanin, s. Alashan, 5—14 Aug. 1908, Koz- low (ZIL); 1 2, Dundusaichan, Gobi, 9 July 1909, Kozlow (ZIL). China. — 1 9, China, 801, 48, curvatum Sm det. Kohl (RMNH); 1 ©, Peking, C. F. Wu coll. (BMNH); 1 ©, Nord Peking, A. David, 1865 . (MNHN); 1 2, “Chasyang”, A. P. Jacot, Aug. 1921 (MNHN); 1 6, Petaiho Beach, Hopei Prov., Kina, 1942, A. M. Hemingston (RMNH); 1 ©, Heishan, Liaching, C. F. Wu coll. (BMNH). Korea. — 3 2, Geumgok, Jinyang-gun, Gyeongnam, July—Aug. 1984, 2 2, Mt. Jiri, 17 July 1984, all G. J. Jeong (RMNH); 1 6, Shoyozan, 30 May 1943, K. Tsuneki (RMNH); 3 ©, Korea, Kim coll., (BMNH); 1 ©, “Corea” (BMNH). USSR. — 18 3 ©, Vladivostok, Sedanka, Ma- laise (NRS); 2 2, Vladivostok, Suchan, Malaise (NRS); 1 2, Ussuri (USNM); 1 2, Okeanskya, Siberia, Aug. 1923, Cockerell (USNM); 1 ®, Suputnskiy zap-k., Primorsky Kraj, Lelej (RMNH) (45° N, 136° E); 1 2, Kongaus, Sibe- ria, Aug. 1923, Cockerell (USNM); 2 ©, Onon, Amour (MNHN); 2 2, Amour, Siberia (BMNH); 1 2, Okeanskaya, Ussuri, 3 July 1911, Cherskiu (ZIL); 1 2, Yakovlevka, Ussuri, 12 Sep. 1926, Diakonov Filipev (ZIL); 1 2, Pri- morje Dist., 20 km E. Ussurijsk, Goznotajozh- noje, 23 July 1983, E. Budrys (ZIL). Sceliphron deforme nipponicum Tsuneki (fig. 48) Sceliphron deforme; Kohl, 1918: 123 (Hokodadi); Tsuneki & Shimoyama, 1963: 48 (Towada dist., Honshu); Tsuneki, 1964: 10 (Mt. Kuju, Sasebo, Kyushu). Sceliphron deforme japonicum Tsuneki, 1971: 7 (2) (nec japonicum Gribodo). Sceliphron deforme nipponicum Tsuneki, 1972: 1, © — Prov. Fukui (Iwaya), paratypes © d, Fukui, R. V. HENSEN: The subgenus Prosceliphron 241 70 71 74 75 76 78 80 Figs. 70—80. Sceliphron rufopictum (Smith). 70, ®, Indonesia, Celebes, clypeus; 71, 3, Philippines, Luzon, cly- peus; 72, S. r. kalshoveni ssp. n., 9, holotype, antennal insertion; 73, S. r. rufopictum (Smith), ©, antennal inser- tion; 74, S. r. kalshoveni ssp. n., 2, holotype, pronotum, lateral aspect; 75, S. r. laticinctum ssp. n., 2, holotype, pronotum, lateral aspect; 76—77, S. r. rufopictum (Smith), ®; 76, first metasomal segment; 77, propodeal ori- fice; 78—80, S. r. laticinctum ssp. n.; 78, 2, Philippines, Luzon, first metasomal segment; 79, 9, Philippines, Luzon, propodeal orifice; 80, 4, paratype, first metasomal segment. 70, 71, 76, 78, 80: scale-line; 74, 75: 2.0 X scale-line; 72, 73, 77, 79: 4.0 X scale-line. 242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) several localities (coll. Tsuneki; not examined); Bohart & Menke, 1976: 106. Description. — Female. — Post-antennal tu- bercles large, but distinctly smaller than in S. d. atripes, and not truncate dorsally; colour pat- tern (fig. 48) more or less like Eastern specimens of S. d. atripes; post-antennal tubercles black, scape dorsally black, subtegular spots small, basal propodeal spots absent; metasoma with the bands more or less darkened (brownish) and reduced; second to fifth sternite black; legs brownish black, with yellow streaks along fore and middle tibiae. The colour-pattern was de- scribed in full detail by Tsuneki (1971, 1972). Male. — Like the female, but darker: clypeus sometimes entirely black, metasoma usually black, except for a narrow brownish band on the first tergite. Material examined. — Japan. — Mt. Haku, 3 9, 1—2 Aug. 1953, 1 d, 1 Aug. 1962; 2 ©, Fu- kui Pr., 8 Aug. 1953; 1 &, Nikko, 3 July 1952; 3 36 3 2, Koike, Fukui, several dates; 2 &, Simou- tinami, Fukui, 30 July 1971; 3 ©, Arashi, Fukui, several dates; 1 2, Ichinose, Mt. Haku, 30 July 1964; 2 2, Hatogaya, Fukui, 29 Aug. 1969; all K. Tsuneki (RMNH); Tamba, 2 ©, Yohoku, 23 July 1955, Higuchi, 1 2, Yakami, 23 July 1955, Honda, 1 & 1 9, Sasayama, 15 July 1955 and 9 Sep. 1951, K. Iwata (all RMNH); 4 9, Japan, Smith coll. (BMNH). Sceliphron fervens (Smith) (figs. 49—60, 117) Pelopoeus fervens Smith, 1858: 101, 2 — Borneo, Sa- rawak, leg. Wallace (OUM; examined). Sceliphron fervens; Turner, 1912: 196; Kohl, 1918: 130, fig. 28. Sceliphron (Prosceliphron) fervens; Bohart & Menke, 1976: 106. Type. — The lectotype, by present desig- nation, is a female in the OUM with only the original Wallace-label “SAR”. The BM pos- sesses three paralectotypes with additional la- bels “P. fervens Smith” in Smith’s handwriting. Two of these have been part of Smith’s own col- lection, according to their label “F. Smith coll., 99—363”, the third is labelled “57—36, Bor- neo . Description. — Body length: 9 15.3—16.5 mm, d 14.5 mm; length of forewing: © 10.6— 11.9 mm, d 9.6 mm. Pubescence greyish white, | tomentum of face silvery. Morphology: Female. — Clypeus with lateral incisions (fig. 49); post-antennal tubercles small (fig. 50); pronotum with median impression; | mesoscutum with sharp transverse striation; laterally with some shallow punctation; propo- deal orifice with narrow dorsal margin, like in S. deforme, but not evenly rounded (fig. 51); pe- tiolus moderately long, distinctly curved (fig. — 57), first tergite distinctly swollen, like in S. de- forme. IFR 0.88—0.95; PTR 0.92—1.00; PR 0.09—0.10; TR 0.26—0.30; SR 0.76—0.82. Male. — In structural characters like the female, but clypeal margin laterally from the small, tri- angular lobes shallowly emarginate (fig. 52), like in S. deforme; petiolus longer and more strongly curved (fig. 55); genitalia similar to those of S. deforme, but volsellar plate more dif- ferentiated and gonostyles apically not so sharp- ly pointed (fig. 53, 54); the difference in the dig- itus is probably an artifact due to dehydration. IFR 0.95; PTR 1.12; PR 0.12; TR 0.29; SR 0.63. Coloration: scapes dorsally black, post-an- tennal tubercles black, upper subtegular spot larger than lower, basal spots of propodeum small or absent, legs mainly reddish, mesosoma often reddish on basal tergites, a distinct yellow band is only present on the third tergite, sometimes the second and fourth bear reduced bands; wings clear, with well-defined dark mark at apex of forewing. Variation: In specimens from Malaysia, Su- matra and Java the red colour on the metasoma is restricted to the first tergite, (fig. 58), whereas in most specimens from Borneo it covers three or more tergites (fig. 59); the one specimen from Palawan I have seen has a pair of spots at the apex of the dorsal enclosure, and the femora are entirely reddish (fig. 60). Distribution. — Southern Thailand, Malaysia, Sumatra, Bangka, W. Java, Borneo, Palawan (fig. 117). Material examined. — Thailand. — 1 ©, Sa- tun, Thaleban Nat. Park, 26 July 1986, R. Hen- sen (CH). Malaysia. — 1 2, Kedah, near Jitra, 8 April 1982, 1 9, Perak, Batang Padang, Jor Camp, 1800 ft., 3 2 1 d, Kuala Lumpur, Gardens, 22 March 1941, all H. M. Pendlebury (BMNH, RMNH); 1 ©, Taiping, W. B. Orme (BMNH); 1 2, Labuang Padang, July 1907, C. B. H. Hunt R. V. HENSEN: The subgenus Prosceliphron (BMNH); 1 2, Kuala Lipis, 29 May 1928, Mill- er (BMNH); Penang, Batu Feringgi, catchment Sree la Basden, 1.9, 14-Reb.1957, 12, 15 Feb. 1961 (BMNH); 1 ?, Ulucheka, in jungle, 10 Aug. 1928, Miller (BMNH). Bangka. — 1 ©, Bangka, Van den Bossche (RMNH). Sumatra. — 1 ©, S. Sumatra, Res. Benkoelen, Boekit-Item, 24 June—2 July, 1935, 650 m, M. E. Walsh (RMNH); 1 2, Muara Sako, Oct. 1915, E. Jacobson (USNM). Java. — 2 2, Radjamandala, Dec. 1938, J. v. d. Vecht (RMNH); 1 2, Radjamandala, Djam- pang Wetan, Oct. 1936, 1200 ft., M. E. Walsh (RMNH); 1 ®, Palaboean Ratoe, 16 April 1933, M. Lieftinck (RMNH). Borneo. — 2 ©, Bettotan, near Sandakan, 11 Aug. 1927 (BMNH); 1 9, E. Borneo, Ketapan, 50 m, June 1937, M. E. Walsh (RMNH); 1 &, Sarawak, Kampong Pueh, Lundu Dist., 690— 1500 m, 25—31 May 1958, T. C. Maa (BPBM); 2, S. E. Borneo, Wahnes S., Wolf v. Schönberg V. (ZMB). Palawan. — 1 ©, 3.2 kmS. of Tarumpitao Pt., 31 May 1958, near jungle ravine, H. E. Milliron (USNM). Sceliphron coromandelicum (Lepeletier) (figs. 61—69, 115) Pelopoeus coromandelicus Lepeletier, 1845: 306, 2 — Coromandel (coll. Spinola, MZU). Sceliphron coromandelicum; Dutt, 1912: 216—221, pl. XII, figs. 1, 7 (bionomics at Pusa, India); Field, 1914: 378—379 (bionomics); Strand, 1915: 91 (Sri Lanka); Kohl, 1918: 125, 126(@ d). Sceliphron (Prosceliphron) coromandelicum; Van der Vecht & Van Breugel, 1968: 192 (type-species of Prosceliphron); Bohart & Menke, 1976: 106. Type. — The Spinola-collection (MT) con- tains one female of this species, standing under the label “Pelopoeus coromandelicus m. et Lep.”, which is most probably the holotype, since the description seems to be based on one specimen. Van der Vecht (pers. comm.) saw the specimen several years ago, and confirmed the general interpretation of the species, on which Kohl (1918) was not sure. The type is in poor condition, the head and mesosoma have been damaged severely by Anthrenus. Description. — Body length: 2 17.4—21.0 mm, d 16.4—19.5 mm; length of forewing: ? 11.9—13.3 mm, d 10.8—11.7 mm. Pubescence: long, erect pubescense of head and mesosoma dark brown or black, but sometimes discolored - 243 to pale brown in older specimens, short pubes- cense as usually white; tomentum of face sil- very. Morphology: Female. — Clypeus with small lateral incisions (fig. 65); post-antennal tuber- cles very small (fig. 63); pronotum with median impression, rather broad in lateral view (fig. 69); mesoscutum strongly transversely striate, only laterally with some shallow punctures; dorsal margin of propodeal orifice narrow (like in S. deforme), but not evenly rounded (fig. 64); petiolus long, sometimes as long as in S. funes- tum, but always distinctly curved (fig. 68). IFR 0.90—0.96; PTR 0.86—0.91; PR 0.05—0.08; TR 0.14—0.18; SR 0.67—0.74. Male. — Like the ©, but clypeus evenly rounded, with small, triangular apical lobes (fig. 66); genitalia (fig. 61, 62): pubescence of cuspis dense, but shorter than in S. rufopictum and S. formosum, volsellar plate reduced, gonostyle with comparatively long and dense pubescence, the lamellae of the gonostyle are rather narrow. IFR 1.00—1.05; PTR 0.97—1.00; PR 0.07—0.08; TR 0.16— 0.20; SR 0.51—0.66. Coloration: Black; the following parts are yellow: two spots on the clypeus (often co- alescent); ventral side of scape, more or less de- veloped band on pronotum (absent in specimens from Malaya), very small subtegular spots; rare- ly a small spot on the scutellum and at apex of the propodeum; petiolus yellow, legs reddish yellow, except coxae, trochanters, base of fore and middle femora and terminal tarsomeres (hind tibiae black in a specimen from Laos). Distribution. — India, Sri Lanka, Bangladesh, Burma, Thailand, Laos, Malaya (fig. 115). S. co- romandelicum is at present the only species known to occur sympatrically with several oth- er Prosceliphron-species: with S. fervens in Ma- laya, with S. deforme femorale in Thailand and Laos, with S. deforme tibiale and S. rectum in Northern India. The ZMA possesses a series of 15 © 2 d from Coimbatore, India, which is almost completely stylopized. Material examined. — Sri Lanka. — 1 9, Pu- wakpitiya, Hiver 1906—7, E. Bugnon (MNHN); 1 2, Mon. Dist., Bibile, 7 June 1975, S. L. Wood & J: L. Petty (USNM); 1 2, Gal Dist., Kanneliya, 24—26 Jan. 1979, 1 2, Tri. Dist, Trincomalee, 0—100 ft, 13—17 May 1976, 2 2, Col. Dist., Labugama Res., 110 m, 29, Oc 1977 and 11-]July1973,1.2,, Rat Wises TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) 244 ‘eIsouopuy “3 WA Jap UEA unpumıg ‘4 °S ‘ZB seaef ‘visauopuy “è “u ‘dss 1uanoqsppg ‘4 °S “Tg “sur ‘$9Q9]9O ‘eIsouopyy “3 (tqarwg) ungsıdofnı ‘4 ‘5 ‘Eg fequing aned Ino]oo ‘(uarus) wnindofns uorgdiars *¢g—]g rie! 245 R. V. Hensen: The subgenus Prosceliphron -1ed 1n0[09 ‘TE1010]N ‘eIsauopyy ‘ VY Ae 98 ö ee eS È € (qus) wnqwrspfiun uosgdija»s ‘Gg € ‘uroned mnojoo ‘puejsuoond) ‘erpemsny ‘à (qarwg) zunsowuso/ [ uougd1120S ‘98 Sura ‘uJoned Ino]oo *adArojoy “è “u ‘dss uunpunngo] wunjndofns uorqdyars ‘pg ‘98—+8 SSA 246 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Figs. 87—95. 87—90. Sceliphron murarium (Smith), 3, Indonesia, Ambon; 87, genitalia, ventral aspect; 88, in- ner side of left half of aedeagus; 89, clypeus; 90, first metasomal segment; 91—93, S. murarium (Smith), ©, In- donesia, Ambon; 91, propodeal orifice; 92, clypeus; 93, first metasomal segment. 94—95, Sceliphron unifascia- tum (Smith), 9, Indonesia, Obi; 94, pronotum, lateral aspect; 95, first metasomal segment. 89, 90, 92, 93, 95; scale-line; 94: 2.0 X scale-line; 91: 4.0 x scale-line; 87, 88: 8.0 x scale-line. R. V. Hensen: The subgenus Prosceliphron 247 Uggalkaltota, 23—26 June 1978, 1 2, Bad. Dist., Ulhitiya, 15 mi. NNE of Mahiyangana, 5—6 Sep. 1980, all leg. K. V. Krombein (USNM); 1 2, Rat. Dist., Udalawaya, 5—6 June 1975, D. H. Messersmith e.a. (USNM); 1 2, Col. Dist., Kollupitiya, 21—22 Nov. 1978, G. Ratnaweera (USNM); 1 ©, Kantalai, 29 June 1953, F. Keiser (RMNH); 1 2, Col Dist., Labu- gama Res., 400 ft., 2—3 Oct. 1976, G. F. Hevel (RMNH); 1 2, Gal. Dist., Kanneliya, 28 July 1973, 300 ft., G. Ekis (RMNH); 1 9, M. Illupa- la, Jan. 1912, Butt.-Repp (ZMB); 1 ©, Bibile, Bad. Dist., 1 2, Ruhuan Nat. Park, 5 Aug. 1963, both Univ. London Ceylon Exp. (BMNH). India. — 3 ©, Nilgiri Hills, Singara, 3400 ft. June 1948, 1 ©, Walayar Forest, 100 ft., 18 Oct. 1947, 1 ©, Kurumbagaram, 12 June 1947, 1 9, Coimbatore, Nov. 1950, all P. S. Nathan (USNM); 1 2, Balasore, R. Oberthür, 1898 (MNHN); 1 2, Mahé, Aug. 1892 (MNHN); 1 2, Ostindien, Dald (ZMB); 4 © 1 gd, Lonavla, W. Ghats, 650 m, 1 à, Sinhagad, near Poona, 1320 m, all May 1963, F. Wain (RMNH); 1 9, Sangli (MBUD); 1 2, Calcutta, 6 June 1944, D. E. Hardy (USNM); 1 ©, Karikal, Nov. 1961, P. S. Nathan (LACM); 1 2, Bangalore, 1894, Bingham (USNM); 7 2, Coimbatore, 1400 ft., 2 2, Anamalai Hills, Chinchona, 3500 ft., 1 ®, Walayar Forest, 700 ft., all April-May 1960, P. S. Nathan (RMNH); 2 ©, Orissa, 11 April 1965, S. D. Jayakar & H. Spurway (RMNH); 1 9, Khasia, 1 ©, Bengalen, 1 ©, Karivar, 6 Aug. 1907 (all RMNH); 1 ©, Bengal (TMA); 15 9 2 3 Coimbatore, 500 m, 7 April 1970, R. T. Si- mon Thomas (ZMA; 2 ® CH); Coimbatore, 495 amy 2) 2), July 1963, 1 2, Nov. 1962, 1 9, April 1964, all P. S. Nathan (ZMA); 2 @, Ka- raikkal, June-July 1964, P. S. Nathan (ZMA); 6 ©, Barrackpore, Rothney (OUM); 5 ©, Bom- bay, W. Ghats, Mathera, Charlotte Lake, 1 April 1908, G. B. Longstaff (OUM); 1 6, India, 5 May 1909, E. Brunetti (BMNH); 2 2, Bom- bay, Matheran or Mableshwar, I. Newton (BMNH); 1 2, Coimbatore, 1925 (BMNH); 1 SAMI hekkadiseWPRenyar Dam Oct. 1938 (BMNH); 1 ©, N. Khasia Hills (BMNH); 2 9, Coimbatore, June 1935, P. S. Nathan (BMNH). Burma. — 10 2 1 d, Rangoon Dist., 1 d, Te- nasserim, Taungoo, all coll. Bingham (ZMB). Thailand. — 1 2, Lee, 22 July 1952, D. & E. Graham (USNM); 1 2, near Metah Valley, J. D. H. Hedley (BMNH). Laos. — 1 ©, Sedone Prov., Pakse (15°10 N, 106°00 E), 15 Aug. 1967, native coll. (RMNH); 1 2, Indochine Francaise, Vitalis de Salveza, Mme A. Vuillet, 1920 (MNHN); 1 2, Vien- tiane, 30 April 1967, native coll. (BPBM). Malaya. — 7 2 4 6, Penang, ex nido, April 1966, H. T. Pagden (BMNH, RMNH); 1 6, with part of nest, from gun barrel, 21 Oct. 1980 (BMNH). Sceliphron rufopictum (Smith) (figs. 70—84, 96—97, 117) Description. — Pubescence: Erect pubes- cence of head and mesosoma yellowish, tomen- tum of face pale-golden. Morphology: Female. — Clypeus with lateral incisions (fig. 70); post-antennal tubercles vari- able in size; vertex sometimes raised; interocu- lar distance at vertex shorter than or equal to length of first flagellomere; pronotal collar vari- able (figs. 74, 75); mesoscutum sharply trans- versely striate, only laterally with shallow punc- tation; propodeal orifice with broad dorsal mar- gin, which is generally brownish translucent (figs. 77, 79); petiolus distinctly curved, shorter than hind tibia, first tergite moderately long, not swollen (figs. 76, 78). Male. — Similar to the female, but clypeus rounded, with small, trian- gular lobes (fig. 71); petiolus longer than hind tibia (fig. 80); genitalia (figs. 96, 97): gonostyles apically rounded, and with broad lamellae, cus- pis with rather dense and long pubescence laterally, volsellar plate rounded triangular. Distribution. — Sunda-islands from Eastern Java to Flores, Celebes, Philippine Islands (fig. 7). Four subspecies can be distinguished, differ- ing in colour pattern as well as in a number of structural characteristics: 1) the size of the post- antennal tubercles, 2) the swelling of the ocellar area, 3) the broadness and median impression of the pronotum. These characters appear to vary clinally from South to North. Recognition of these clines led me to regard the geographic forms as conspecific. No conclusions could be drawn out of male genital structure, since only one male specimen was available. Key to the subspecies of Sceliphron rufopictum (Smith) 1. Post-antennal tubercles rather large (fig. 73), sometimes with yellow spot; subtegu- lar and basal propodeal spots large (fig. 83, Ban ben th Basar Alea, 2 — Post-antennal tubercles small (fig. TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) 248 ‘39adse [enusA “eıferrusd ‘66 ;sn3eapae Jo Jeu 279] FO APIs JOUUT ‘86 ‘JEULOTEPEND) ‘spue[s] UOWOJOS “Pp ‘[UOM 24977390 unsowsof uou -441 295 ‘66—86 ‘sndvapar zo Fey 159] Jo apts Jouur ‘76 fivadse jenuoa “eıferuag ‘96 SadAreıed ‘9 “u “dss wnqouiniy) wnsıdofnı uoLgdyars ‘L6—96 '66—-96 SBH 66 86 16 um Q°T R. V. Hensen: The subgenus Prosceliphron 249 black; subtegular and basal propodeal spots Sn ASE veer cue EL PRE ee. 3 2. Vertex raised behind anterior ocellus; usually tergites 2—5 with complete yellow band, antennal scape dorsally black; Phil- ippine Islands +... 42"... laticinctum ssp. n. — Vertex not or hardly raised; only third ter- gite with distinct apical band, antennal Scapemeddish dorsally; Gelebes............ EN. iI rufopictum (Smith) 3. Fifth tergite with yellow band, base of fem- ora black (Java) or femora entirely reddish (Hors) N tn... kalshoveni ssp. n. — Fifth tergite black, femora totally reddish; Sumbat. ke bicinctum Van der Vecht Sceliphron rufopictum kalshoveni ssp. n. (figs. 72, 74, 81, 117) Holotype. — ©, M. Java, S. Coast, Patjitan, 12 Dec. 1937, J. van der Vecht (RMNH). Description. — Body length 16.6 mm, length of forewing 10.7 mm. Morphology. — Post-antennal tubercles small (fig. 72); ocellar area hardly raised; prono- tal collar narrow (fig. 74), with distinct median impression; propodeal orifice rounded, upper margin very broad (cf. fig. 77). Coloration. — Black; the following parts are yellow: broad spot on clypeus, antennal scapes, dorsal band on pronotum, spots on the tegulae, small subtegular spots, transverse spot on scu- tellum, very small basal propodeal spots, trian- gular mark at apex of propodeum, first metaso- mal tergite, and bands on tergites 2—5; legs light reddish, base of femora black, yellow spots at apex of fore and middle femora, yellow lines anteriorly on fore and middle tibiae (fig. 81). Paratypes. — Java. — 1 2, Gedangan, 8 Sep. 1939, P. A. Blijdorp; 2 2, Gedangan, 18 Feb. 1933, Kalshoven; 1 9, Djati Rogo, Djati Forest, 100 m, M. E. Walsh; 1 2, Midden Java, with nest, Kalshoven (all RMNH). Karimun Djawa Islands. — 2 9, 22—30 Nov. 1930, M. A. Lieftinck. Not paratypes: Flores. — 2 9, W. Flores, Pater Verheyen, no. 116 (RMNH). Variation. — Body length 16.6—18.8 mm, length of fore wing 10.8—12.3 mm; IFR 0.89— 0.95; PTR 0.82—0.90; PR 0.04—0.07; TR 0.13—0.19; SR. 0.76—0.84. The specimens from Flores have the femora entirely reddish, and the basal propodeal spots and the subtegu- lar spots are comparatively large. These speci- mens are probably transitional between the sub- species S. r. rufopictum, r. kalshoveni, and r. bi- cinctum. Sceliphron rufopictum bicinctum Van der Vecht (figs. 82, 117) Sceliphron deforme bicinctum Van der Vecht, 1957: 370, 2 — Sumba (Museum Basel; not examined). Sceliphron (Prosceliphron) deforme bicinctum; Bohart & Menke, 1976: 106. Description. — Body length 17.4—18.0 mm, length of forewing 11.0—11.2 mm. Morphology. — Post-antennal tubercles small (cf. fig. 72); vertex not raised; pronotal collar with distinct median impression (cf. fig. 74); propodeal orifice rounded, with broad dor- sal margin (cf. fig. 77). IFR 0.93—0.95; PTR 0.85—0.87; PR 0.07; TR 0.16—0.20; SR 0.71— 075% Coloration. — Post-antennal tubercles black, antennal scapes yellow, subtegular spots and basal propodeal spots small, femora, tibiae and tarsi reddish, first tergite reddish, third and fourth tergite with yellow apical band (fig. 82). Material examined. — Sumba. — 1 9, W. Sumba, Pogobina, 16 Sep. 1949, 1 2, O. Sumba, Laluku, 4—7 July 1949, both leg. Dr Buhler & Dr Sutter (paratypes, RMNH). Sceliphron rufopictum rufopictum (Smith) (figs. 73, 76—77, 83) Pelopoeus rufopictus Smith, 1856: 232, 2 — Celebes (BMNH; examined). Pelopoeus flavo-fasciatus Smith, 1859: 15, 2 — Ce- lebes, leg. Wallace (OUM; examined). Sceliphron rufopictum; Kohl, 1918: 129. Sceliphron deforme rufopictum; Van der Vecht, 1957: 370. Sceliphron (Prosceliphron) deforme rufopictum; Bo- hart & Menke, 1976: 106. Types. — The lectotype of P. rufopictus Smith, by present designation, is a female la- belled “Celebes, 55/22” in the BMNH, which is the only specimen old enough, according to la- bel data, and which fits the original description. Previously a specimen with label “China, 56/45”, belonging to S. d. deforme (Smith), was erroneously marked as the type. The lectotype of P. flavofasciatus Smith, by present designation, is a female with labels 250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) 113 112 1.0 mm ____i R. V. HENSEN: The subgenus Prosceliphron 251 Fig. 114. Distribution of (1) Sceliphron shestakovi Gussakovskij, (2) Sceliphron funestum Kohl, (3) Sceliphron rectum pulchellum Gussakovskij, and (4) Sceliphron r. rectum Kohl. “Celebes” and “Pelopoeus flavo-fasciatus Smith” (Smith’s handwriting) in the OUM. Par- alectotypes are 2 2 with labels “Mak” (Makas- sar) (Wallace’s handwriting) and “Pelopoeus fla- vofasciatus Smith” (Smith’s handwriting) (BMNH and OUM). The specimen in the BMNH is additionally labelled “F. Smith coll., BM 79—22” and is stylopized. Description. — Body length 17.8—21.6 mm, length of forewing 11.4—13.7 mm. Morphology. — Post-antennal tubercles rath- er large; vertex sometimes slightly raised behind anterior ocellus; pronotal collar with distinct median impression, rather broad in lateral view (fig. 75); dorsal margin of propodeal orifice rounded and broad (cf. fig. 77). IFR 0.84—0.96; PTR 0.76—0.85; PR 0.05—0.08; TR 0.14— 0.18; SR 0.80—0.86. Coloration. — Antennal scapes yellow; post- antennal tubercles usually with yellow spot; Figs. 100—113. Sceliphron formosum (Smith). 100—101, S. f. ocellare Kohl, 2, Solomon Islands; 100, upper part of head; 101, antennal insertion; 102, S. f. formosum (Smith), ©, Australia, Queensland, antennal insertion; 103—106, clypeus; 103, S. f. formosum (Smith), ®, Australia, Queensland; 104, S. f. bruinjnu (Maindron), ®, N.W. Nw. Guinea; 105, S. f. formosum (Smith), d. Australia, Queensland; 106, S. f. ocellare, &, Solomon Is- lands; 107—109, propodeal orifice; 107, S. f. formosum, ®, Australia, Queensland; 108, °, S. f. ocellare (Kohl), 2, Solomon Islands; 109, S. f. bruinjnit (Maindron), 9, Indonesia, New Guinea; 110—112, first metasomal seg- ment; 110, S. f. formosum (Smith), 2, Australia, Queensland; 111, 112, S. f. ocellare Kohl, 3, Solomon Islands, 111, Nygela, 112, Guadalcanal; 113, S. f. ocellare Kohl, 2, Solomon Islands, Malaita, pronotum, lateral aspect. 103—106, 110—112, scale-line; 100: 0.5 X scale-line; 113: 2.0 X scale-line; 101, 102, 107—109: 4.0 X scale-line. 252 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Fig. 115. Distribution of (5) Sceliphron curvatum (Smith), and (12) Sceliphron coromandelicum (Lepeletier). ' subtegular and basal propodeal spots large; dor- sal enclosure of propodeum sometimes with a pair of spots (N. Celebes); first tergite reddish, third with a broad yellow band apically; femo- ra, tibiae and tarsi entirely reddish (fig. 83). Material examined. — Celebes. — 1 2, Bua Kraeng, 5000 ft., Feb. 1896, 1 ?, Samanga, Nov. 1895, both H. Frühstorfer (NMW); 1 2, Patunuang, Jan. 1896, H. Frühstorfer (ZMB); 1 ©, Toelabella, Rosenberg (RMNH); 1 9, Lom- poh Batang, 200 m, 1941, H. Lucht (RMNH); 2 ©, Manado, June and Aug. 1941, F. Dupont (RMNH); Bantimurung, 1 2, 3 Oct. 1931, J. van der Vecht, 1 ©, July 1949, C. J. H. Frans- sen, 1 2, G. Ribbe, 1882 (all RMNH); 1 9, Lo- ka, 1200 m, C. J. H. Franssen (RMNH). Sceliphron rufopictum laticinctum ssp. n. (figs. 75, 78—80, 84, 96—97) Sceliphron deformis; Williams, 1919: 122, fig. 59 (Lu- zon, bionomics); Rohwer, 1921: 675 (Luzon). Holotype. — 2, Los Banos, Philippine Isl., July-Aug. 1917, F. X. Williams coll. (RMNH). Description. — Body length 18.3 mm, length of forewing 12.0 mm. Morphology. — Post-antennal tubercles rath- er large, shiny (fig. 73); vertex moderately raised behind anterior ocellus; pronotal collar evenly rounded, without traces of a median im- pression, rather broad in lateral view (fig. 75); mesosoma more regularly and finely sculptured than in the other subspecies, and more shiny, particularly the mesopleuron; propodeal orifice more or less trapezoid in outline, dorsal margin broad and brownish translucent (fig. 79). Coloration. — Black, the following parts yel- low: large spot on clypeus, scapes ventrally, broad band on pronotum, large transverse spot on scutellum, large subtegular spots, small spots on tegulae, basal spots and apical spot on pro- podeum, first metasomal tergite, except on the middle, apical bands on tergites 2—5 (the one on the second tergite less strong), faint bands on sternites 2—5; reddish are all tibiae, tarsi, and the apical 2/5 of the femora, but the last tarso- meres are infuscated, and the fore and middle tibiae bear a yellow stripe anteriorly. The col- our-pattern is illustrated in fig. 84. Measurements (entire type-series). — Body length 16.0—20.0 mm, length of forewing 10.8—13.0 mm; IFR 0.82—0.87; PTR 0.80— 0.89; PR 0.08—0.09; TR 0.17—0.23; SR 0.76— 0.88. Male. — Body length 15.8 mm, length of forewing 10.6 mm. IFR 0.81; PTR 1.02; PR R. V. HENSEN: The subgenus Prosceliphron 253 Fig. 116. Distribution of the subspecies of Sceliphron deforme (Smith): (6) S. d. tibiale Cameron, (7) S. d. femo- rale ssp. n., (8) S. d. deforme (Smith), (9) S. d. atripes (Morawitz), and (10) S. d. nipponicum Tsuneki. 0.10; TR 0.17; SR 0.60. Morphologically like the 2, coloration see below (specimen from Ce- bu). Paratypes. — Luzon. — Los Banos, 1 ?, 23 Sep MOIS AVS Sur (iO 920 July ZINE: Manuel, 2 2, 7 Sept. and 8 Nov. 1953, Townes Family, 1 2, 30 May 1954, H. & M. Townes, (all RMNH); 1 2, Limay, Bataan, Sep. 1920, 1 ©, Camalia, Albay, 26 Aug. 1893, A. E. Bigor- nia, 1 2, Mt. Maquiling, 3 Feb. 1954, M. Delfi- nado, 1 ®, Altimonan, Quezon, July 1963, A. Concepcion (all RMNH); 1 ©, Montalban, Baker, 2 2, Mt. Maquiling, Baker (all USNM). The following specimens are not included in the type-series. They originate from other is- lands of the Philippine Archipelago, and exhibit constant differences in colour-pattern. The specimens from Mindanao may even be re- garded as transitional between S. r. rufopictum and S. r. laticinctum. Samar. — 5 Q, Island Samar, Baker (USNM). (With more or less distinct median impression on pronotum; antennal scape with narrow lon- gitudinal black line dorsally; femora to a larger extent reddish). Cebu. — 1 2 1 d, Camp 7.25 km W. Cebu City, 21—29 Sep. 1965, D. Davis, 400 m 254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) | (USNM). (Pronotal collar with median impres- sion, antennal scape with black line dorsally, only third tergite with distinct apical band; first and second tergite of male partly reddish, fore and middle tibiae of male posteriorly black). Bilirany MSP Islande Biliran, Baker (USNM). (Hind tibiae reddish, broad band on second tergite). Mindanao. — 4 ©, Dapitan, Baker (USNM); 12077 Dayao, April 1027 Vie ne. Greco (BMNH). (Distinct median impression on pro- notum, large yellow spots on fore and middle femora and large spots on dorsum of propo- deum (like S. r. rufopictum-specimens from North Celebes)). Sceliphron murarium (Smith) (figs. 87—93, 117) Pelopoeus murarius Smith, 1863: 34, © — Ceram, leg. Wallace (OUM; examined). Pelopoeus rufipes Mocsary, 1883: 24, 2 — Ambon (TMA; not examined) (nec rufipes Fabricius, 1804). Sceliphron mocsaryı Dalla Torre, 1894: 388 (new name for P. rufipes Mocsary). Sceliphron rufipes; Kohl, 1918: 131. Sceliphron (Prosceliphron) fervens murarium; Bohart & Menke, 1976: 106. Type. — In the Saunders-collection (OUM) two specimens are standing under this species. One of these is a female with labels “Cer.” and “Pelopoeus murarius Smith” (Smith’s hand- writing). This specimen is the lectotype by pre- sent designation. The other specimen belongs to S. formosum bruinjnu (Maindron) and is a male, labelled “N.” and “Pelopoeus murarius Smith male?” (Smith’s handwriting). It was evidently not part of the original type-series. Description. — Body length: 2 17.8—21.0 mm, d 16.8—17.7 mm; length of forewing: © 12.5—14.0 mm, d 11.5—12.1 mm. Pubescence: erect pubescence of head and mesosoma dark brown; tomentum of face golden. Morphology: Female. — Clypeus with lateral incisions (fig. 92); post-antennal tubercles very small (cf. fig. 50); vertex not raised; pronotum with distinct median impression, high and nar- row in lateral view, like in S. unifasciatum (fig. 94); mesonotum sharply transversely striate, only laterally with a trace of punctation; propo- deal orifice nearly circular in outline, dorsal margin very broad (fig. 91), and brownish translucent; petiolus rather short (shorter than in S. unifasciatum and S. rufopictum), first ter- gite long but not swollen (fig. 93). IFR 0.81— 0.88; PTR 0.71—0.75; PR 0.06—0.07; TR 0.15—0.22; SR 0.80—0.95. Male. — In most re- spects like the female, but clypeus rounded, with small triangular lobes (fig. 89); petiolus | long, strongly curved (fig. 90); genitalia (fig. 87, | 88) like those of S. rufopictum, but more stoutly built, with shorter aedeagus and gonostyles, and more strongly curved digitus, however with the same dense flock of setae laterally on the cuspis. IFR 0.89—0.93; PTR 0.83—0.89; PR 0.08— 0.09; TR 0.16—0.17; SR 0.65—0.70. Coloration. — Sufficiently described by Kohl (1918) after the type of S. rufipes Mocsary. Most important traits are the absence of yellow mark- ings on the mesosoma, except for occasional traces of a band on the pronotum; the first and second tergite and the second sternite are red- dish, the remaining tergites more or less fuscous apically. Distribution. — Endemic to the Southern Moluccas (fig. 117). Material examined. — Ambon. — 1 2, Am- bon (RMNH); 4 ©, Ambon, Waai, 1—50 m, 12—26 May 1965, 2 2,1 6, Ambon Isl., 70 m, resp. 10 and 26 Nov. and 23 Sep. 1960, 3 9, Ambon, 70 m, resp. 22 Jan., 1 April and 7 May 1961, all A. M. R. Wegner (RMNH); 10 9 1 6, Ambon, Waai, different dates, A. M. R. Wegner (BPBM). Ceram. — 1 ®, West Ceram, April—June 1910, Van Dalen (ZMA). Sceliphron unifasciatum (Smith) (figs. 85, 9495, 117) Pelopoeus unifasciatus Smith, 1861: 123, 2 — Batjan, leg. Wallace (OUM; examined); Kohl, 1918: 129 (in synonymy of S. rufopictum). Pelopoeus affinis; Maindron, 1878: 395, pl. IX, fig. 9, © — Halmaheira (MNHN) (nec Sphex affinis Fabricius, 1793); Kohl, 1918: 129 (doubtful syno- nym of S. rufopictum), 132 (original description). Sceliphron (Prosceliphron) deforme unifasciatum; Bo- hart & Menke, 1976: 106. Type. — There is one specimen of this species in the Saunders-collection (OUM), labelled “Bac” (Batjan) and “Pelopoeus unifasciatus Smith” (Smith’s handwriting), which I regard as the holotype. Description. — Body length 18.2—19.5 mm; length of forewing 12.1—13.1 mm. Pubescence: erect pubescense of head and mesosoma purely R. V. HENSEN: The subgenus Prosceliphron 255 Fig. 117. Distribution of (11) Sceliphron fervens (Smith), (13) S. rufopictum kalshoveni ssp. n., (14) S. r. bicinc- tum Van der Vecht, (15) S. r. rufopictum (Smith), (16) S. r. laticinctum ssp. n., (17) S. murarium (Smith) and (18) S. unifasciatum (Smith). white, not yellowish as in S. rufopictum; facial tomentum silvery. Morphology. — Clypeus with lateral inci- sions (cf. fig. 92); post-antennal tubercles very small; vertex not raised; pronotal collar high and narrow in lateral view (fig. 94), medially impressed; mesoscutum sharply transversely striate, laterally with shallow punctation; pro- podeal orifice more or less rectangular, dorsal margin very broad (like fig. 91); petiolus dis- tinctly curved, longer than in S. murarium, first tergite rather long, but not swollen (fig. 95). IFR 0.88—0.90; PTR 0.72—0.78; PR 0.07— 0.08; TR 0.17—0.21; SR 0.85—0.93. Coloration. — Clypeus black or with small yellow spot; pronotum usually with narrow, in- terrupted yellow band; small subtegular spots present, and usually a small spot on the scutel- lum; metasoma sometimes with central yellow spot on the first tergite, and usually with band on the third tergite. There is considerable varia- tion in colour-pattern between the different is- land-populations; the colour-pattern of a speci- men from Morotai is illustrated in fig. 85. The & of this species is unknown. Distribution. — Endemic to the Northern Moluccas (fig. 117). Material examined. — Halmaheira. — 1 9, Kao Dist., Kampung Toliwang, 1—14 March 1981, A. C. Messer & P. M. Taylor (USNM) (scutellum only with very small spot, first ter- gite black). Morotai. — 2 ®, Morotai, Bernstein (RMNH) (pronotum black, first tergite black (fig. 85)). Obi. — 2 2, W. Obi, Lake Riv., 0—50 m, Ju- ly—Nov. 1953, A. Wegner (RMNH) (prono- tum black, scutellum black, all tergites black). 256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Sceliphron formosum (Smith) (figs. 86, 98—113, 118) Description. — Body length: 2 15.4—20.7 mm, 6 15.2—18.7 mm; length of forewing: 9 10.8—12.7 mm, 6 11.0—12.3 mm. Pubescence: erect pubescense of head and mesosoma yellow- ish, facial tomentum golden. Morphology: Female. — Clypeus with more or less distinct lateral incisions (figs. 103, 104); post-antennal tubercles variable in size; vertex strongly raised in one subspecies; pronotal col- lar with median impression, which may be very shallow; mesoscutum transversely striate, but less sharply than in other species, except S. cur- vatum, moreover with distinct punctation be- tween the striae; propodeal orifice with rather broad dorsal margin (figs. 107—109), which is often yellowish or brownish translucent; petio- lus distinctly curved, shorter than hind tibia, first tergite short and slender (fig. 110). IFR 0.87—1.05; PTR 0.76—0.88; PR 0.06—0.08; TR 0.15—0.20; SR 0.71—0.88. Male. — Like the female, but clypeus rounded, with hardly a trace of lobes, medially emarginate (figs. 105, 106); petiolus about as long as the hind tibia (figs. 111, 112); genitalia (figs. 98, 99): gono- styles pointed, digitus comparatively long and slender, volsellar plate distinctly triangular. IFR 0.93—1.05; PTR 0.88—1.02; PR 0.08—0.10; TR 0.12—0.18; SR 0.55—0.76. Distribution. — New Guinea, Bismarck-Ar- chipelago, Solomon Islands, Northern and East- ern Australia (fig. 118). This species consists of three distinct geo- graphic forms, which differ in colour as well as in some structural details; the large area in which intergradation occurs, however, proves that these forms do not merit specific status. The transitional specimens (from Southern and Eastern New Guinea) are provisionally divided over the subspecies. Key to the subspecies of Sceliphron formosum (Smith) 1. Second tergite black, or with strongly re- duced band; usually the sternites are black, the petiolus and the antennal scape yellow; Australia, SsNewiGuinean ve Nn formosum (Smith) — All tergites with complete yellow bands; sternites with yellow bands or lateral spots; petiolus usually black; scape dorsally black 2. Vertex raised (fig. 100); post-antennal tu- bercles and tegulae with yellow spot; only first tergite partly red; Bismarck-Archipela- go, Solomon-Islands, E. New Guinea...... NAIL ue On oe babies pes ocellare Kohl — Vertex not raised; post-antennal tubercles and tegulae black; often all tergites with reddish ground-colour; N. and W. New Guinea bruinjnii (Maindron) Sceliphron formosum bruinjnii (Maindron) (figs. 104, 109, 118) Pelopoeus bruinjnu Maindron, 1878: 394, pl. 9: 10, 2 — Manokwari (“Dorey”) and Anday, New Gui- nea (MNHN;; examined). Sceliphron bruinjnu; Cameron, 1906: 221 (Merauke, Etna Bay); Kohl, 1918: 134. Sceliphron bruyni Cameron, 1906: 56 (emendation). Sceliphron (Pelopoeus) fallax Kohl, 1918: 129, 26 — Stephansort, Astrolabe Bay, New Guinea (NMW; examined). Sceliphron (Prosceliphron) bruijnü; Van der Vecht & Van Breugel, 1968: figs. 2—4 (genitalia of 3). Sceliphron (Prosceliphron) bruinjnit; Bohart & Menke, 1976: 106. Maindron dedicated this form to Mr. “Bruinjn” at Ternate, whose actual name was “De Bruyn”. Cameron’s emendation however is incorrect, since in the original publication there is no evidence of this error. Types. — The Paris Museum possesses the two syntypes of P. bruinjnü. The female with lables “Dorey” and “Raffray & Maindron, Do- rey, Nouvelle Guinée” is herewith designated as the lectotype. The paralectotype is labelled “Nouvelle Guinée, Andaie, Mai 1878”. The Vienna Museum possesses three syntypes of S. fallax. The lectotype, by present desig- nation, is a female with labels “N. Guinea, Biro 97”, “Stephansort, Astrolabe B.” and “Sc. fallax Kohl”. The male paralectotype is labelled equal- ly; the female paralectotype lacks Kohl’s identi- fication-label. Description. — Morphology. — Vertex not or slightly raised; post-antennal tubercles rather large (cf. fig. 101); propodeal orifice often dis- tinctly trapezoidal in outline, with the dorsal margin straight (fig. 109). Coloration. — Rather variable; the type ıs a brightly coloured specimen, with large subtegu- lar spots, basal propodeal spots and broad bands on the tergites and sternites; the first and the second tergites are almost entirely reddish. Oth- er specimens, like the type-series of S. fallax, are R. V. HENSEN: The subgenus Prosceliphron 257 Fig. 118. Distribution of the subspecies of Sceliphron formosum (Smith): ( f. formosum (Smith), and (21) S. f. ocellare Kohl. darker, without subtegular spots, without red parts on the metasoma, and with the bands on the tergites and sternites very narrow. Material examined. — New Guinea. — 1 9, Manokwari, 23 May 1903 (ZMA); 1 2, Pionier- bivak, July 1920, W. C. van Heurn, 1 ©, Hat- tam, Utrechts Zendingsgenootschap, 1 2, Tami River, Hollandia, 1920, R. Voorhoeve (all RMNH); Ifar, 2 9, 1 March 1957, 1 2 2 6,5 DESSERT TOE MOSER March 1959, all J. van den Assem (RMNH); 1 2, Humboldt Bay, Hollandia, April 1936, L. E. Cheesman, 1 ®, Hollandia, Jan. 1937, 300—600 m (BMNH); 1 ®, Fiume, Purari, Loria, Jan. 1894 (Mus. Genoa); 1 2, Dorey, Humboldt Bay, 1906, O. K. Pasteur (MNHN); 1 2, Me- dang Dist., Wanuma, 600-720 m, Aug. 1968, J. Sedlacek (BPBM). The following specimens are transitional be- tween this subspecies and S. formosum ocellare: S. f. bruinjnii (Maindron), (20) S. New Guinea. — 1 ©, Kwamkwam, 90 km NW. Finschhaten, 18 Jan. 1973, K. W. Ströder (RMNH); 1 2, Wau, Morobe Dist., 1150 m, 17 May 1962, J. Sedlacek (BPBM): these specimens have the vertex slightly raised, the post-antennal tubercles are rather large and marked with yel- low, the tegulae bear very small yellow spots, the metasoma however is largely red; 12, Wau, 1200—1400 m, 18—26 Sep. 1972, J. van der Vecht (RMNH); 1 ©, Hohola, Pt. Moresby, Central Dist., 30 May 1966, in ornamental gar- den (RMNH); 1 6, Port Moresby, emerged from mud-nest (Australian Museum): these specimens have the vertex slightly raised, the post-antennal tubercles are large, and the tegu- lae bear small yellow spots; 1 2, Bulolo, 700 m, 6 Nov. 1969, J. Sedlacek (BPBM); 1 ©, Sim- bang, Huon Golf, Bird, 1900 (Sc. ocellare, type, det. Kohl) (NMW): like the previous, but with additional yellow spot on the post-antennal tu- bercles. 258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Sceliphron formosum formosum (Smith) (figs. 86, 102—103, 105, 107, 110, 118) Pelopoeus formosus Smith, 1856: 230, ® — Australia (BMNH; examined). Sceliphron papuanum Cameron, 1906: 221, 9 — New Guinea, Merauke (ZMA; examined). Sceliphron (Pelopoeus) formosus; Kohl, 1918: 126— 128 (2 3), 138 (original description S. papua- num). Sceliprhon (Prosceliphron) formosum; Bohart & Men- ke, 1976: 106. Type. — The lectotype, by present desig- nation, is a female with labels “Port Essington” and “formosus Sm., type” in Smith’s hand- writing. Another specimen, labelled “type, F. Smith coll. 79—22” is probably not a type-spec- imen: it does not agree with the description, and may be of a later date. Description. — Morphology. — Typically the post-antennal tubercles are small (fig. 102), the vertex is not raised, and the clypeus has dis- tinct lateral incisions in the female (fig. 103). Coloration. — As described by Kohl (1918); the characteristic traits are: post-antennal tuber- cles black, pronotal band not interrupted, pro- podeum with yellow spots at apex of dorsal en- closure, petiolus yellow, second and usually third tergite black, sternites 2—5 black (fig. 86). Variation. — In some N. Australian speci- mens the petiolus is black, and the sternites bear yellow lateral spots; the spots at apex of the dorsal enclosure may be absent; in the extreme North of Australia the morphology tends to be transitional between formosum and ocellare: the post-antennal tubercles are larger and the vertex is slightly raised. Distribution. — Southern New Guinea, East- ern parts of Australia (fig. 118). The occurrence in Tasmania is doubtful. Kohl (1918) mentions also Ceram and Ternate, without reference to specimens or literature, but this is almost cer- tainly incorrect. Material examined. — Australia. — 1 9, S. E. Queensland, 14 Feb. 1957, A. R. Steginga (ZMA); 2 ®, Australia, Feb. 1847, Vereaux (MNHN); 3 2, Tasmania, 1846, Vereaux (MNHN); 1 2, Nov. Hollande, Gory (MINEIN) PMR OE eb 1900 RAME Rurner. (BMNH); 1 2, Stradford, Barron River, 17 Jan. 1972, 1 ©, Cairns, 30 Jan. 1972, both L. Ooster- weghel (RMNH): 1 9%, Australia boreal (RMNH); 1 2, N. S. Wales (RMNH); 2 ©, Brisbane, March 1944, E. F. Riek (USNM); 2 | ?, Stradbroke Island, 5 Dec. 1913, 1 2, Bris- bane, 21 Dec. 1912, both E. F. Riek (USNM); 1 ©, Halifax, Feb. 1913, Girault, 1 ©, Cairns, 1 ®, Sunnybank, 1 Dec. 1951, E. F. Riek (all USNM); 15 d 10 2, Mackay, Jan. 1901, R. E. Turner (BMNH); 1 2, Townsville, 1 March | 1902, F. P. Dodd (BMNH); 2 ©, Tamworth, 25 Jan. 1960, M. Edwards (BMNH); 2 @, Elalie, S. of Elaggy, 9 Nov. 1917, L. Oosterweghel, 1 2, Terra van Diemeni, Parzudaki, 1 ©, Nov. Holl. (all RMNH); 1 ©, Claudie R., 5 Mls. W. Mt. Lomond, Queensland, 23 Dec. 1971, DIR} McAlpine (USNM). New Guinea. — 1 9, New Guinea Exp. 1905/6, Merauke (ZMA; holotype of S. papua- num Cam.); 1 2, Eramboe, 80 km ex Merauke, 5 Feb. 1960, T. C. Maa (BPBM); 1 2, Pt. More- sby area, May 1947, L. Jones (BMNH). Sceliphron formosum ocellare Kohl (figs. 100—101, 106, 108, 111—113, 118) Sceliphron (Pelopoeus) ocellare Kohl, 1918: 128, 2 — Bismarck-Archipelago, Ralum (? ZMB) and Kini- gunang (? NMW). Sceliphron (Prosceliphron) ocellare; Bohart & Menke, 1976: 106. Type. — I have seen a specimen labelled by Kohl as “type”, but it originates from New Guinea, and thus cannot be considered a type- specimen. No other type-material could be found in the NMW, and neither in the ZMB and TMA. Description. — Morphology. — Lateral inci- sions in clypeus of female usually less developed than in other subspecies; post-antennal tuber- cles rather large (fig. 101); vertex moderately to strongly raised behind anterior ocellus (fig. 100). Coloration. — As described by Kohl (1918: 128); the diagnostic features are: antennal scape dorsally black, post-antennal tubercles with yel- low mark, tegulae with yellow mark, petiolus usually black, tergites with broad bands, ster- nites with bands or lateral spots. Variation. — The petiolus and scapes are completely yellow in two males from Manus, which also have the legs nearly entirely yellow; one of these also had a pair of spots at the apex of the dorsal propodeal enclosure. The vertex 1s. extremely swollen in four females from Nor- manby Island; the coloration is very rich in these specimens, including additional spots R. V. Hensen: The subgenus Prosceliphron 259 laterally on the pronotum, on the hypo-epime- ral area, and sometimes laterally on the mesos- cutum. Material examined. — Admiralty Islands. — 2 2, Manus, Lorengau, 17 and 19 June 1962, 2 2, Lavongai, Banatam, 19 and 23 March 1962, all Noona Dan Exp. (UZM). New Britain. — 1 dg, Rabaul, 1933, J. L. Froggatt (BMNH). New Ireland. — 1 9, Kandan, 25 Dec. 1959, W. W. Brandt (BPBM); 1 d, Lemkamin, 16 April 1962, Noona Dan Exp. (UZM). Normanby Island. — 4 ©, Wakaiuna, Sewa Bay, 1—8 Jan. 1957, W. W. Brandt (BPBM, RMNH). New Guinea. — transitional specimens, see under S. f. bruinjnit. Solomon Islands. — 1 ?, Small Gela, Balaga, 12731 Dec: 1963, M. J. A. de Korter, 1 ©, Guadalcanal, Honiara, Nov. 1967 (both ZMA); 1 2, Iles Salom, 1914, E. André (MNHN); 2 2, Malaita, Dala, 50 m, 6—8 June 1964, J. & M. Sedlacek (BPBM); 1 2, Isabel, Rasa, 11 Dec. 1964, M. McQuillan, 5 2 3 &, Guadalcanal, Kukum, 7 Dec. 1961, M. Greenslade, 1 2, Bou- gainville, Oct. 1960, R. W. Paine (all BMNH); 1 2, Tulagi, jungle, collecting mud, 16 Dec. 1934, 1 2, Tulagi, Sasapi cutting, 25 Dec. 1934, 1 9, Sevo Isl., Reko, 23 Feb. 1934, all H. T. Pagden (RMNH); 1 9, Bougainville, Kieta, 26 Nov. 1959, T. C. Maa (BPBM). REFERENCES Basil-Edwards, S., 1921. On the habits of a Sceliphron wasp (S. deforme). — Journal of the Bombay Nat- ural History Society 28: 293—297, figs. 1, 2. Beaumont, J. de, 1969. Beitrage zur Kenntnis der Fauna Afghanistans. Sphecidae, Hymenoptera. — Casopis Moravského Musea, Brno 54 (suppl.): 385—406, 2 figs. Bingham, C. T., 1897. The fauna of British India. Hymenoptera 1: i—xiii + 1—579. — Taylor and Francis, London. Bohart, R. M., & A. S. Menke, 1976. Sphecid wasps of the world, a generic revision: i—ix + 1—695, figs. 1—190. — University of California Press, Berke- ley. Cameron, P., 1899. Description of a new genus and some new species of fossorial Hymenoptera from the Oriental Zoological Region. — Annals and Magazine of Natural History (7)4: 52—69. Cameron, P., 1900. Descriptions of new genera and new species from the Oriental Zoological Region. — Annals and Magazine of Natural History (7)5: 17—41. Cameron, P., 1906. Hymenoptera of the Dutch Expe- dition to New Guinea in 1904 and 1905. Part I. Thynnidae, Scoliidae, Pompilidae, Sphegidae and Vespidae. — Tijdschrift voor Entomologie 49: 215—233. Dalla Torre, C. G. de, 1897. Catalogus Hymenopte- rorum hucusque descriptorum systematicus et synonymicus 8, Fossores: i—viii + 1—749. — G. Engelmann, Lipsiae. Dutt, G. R., 1912. Life histories of Indian insects IV. — Memoirs of the Department of Agriculture in India 4: 183—267, pls. 1—4. Eady, R. D., 1968. Some illustrations of microsculp- ture in the Hymenoptera. — Proceedings of the Royal Entomological Society of London (A) 43 (4—6): 66—72, figs. 1—31. Field, F., 1918. Habits of the Mason-Wasp. — Journal of the Bombay Natural History Society 23: 378— 379. Gussakovskij, V. V., 1928. Sphecidarum Species no- vae. — Trudy Zoologicheskogo Instituta, Akade- miya Nauk SSSR, Leningrad 4: 3—19. Gussakovskij, V. V., 1933. Sphecidae et Psammocha- ridae a cl. N. Zarudnyi in Persia orientali collec- tae. — Travaux de l’Institut Zoologique de l’A- cadémie des Sciences de l'URSS 1: 269—304. Gussakovskij, V. V., 1936. Schwedisch-chinesische wissenschaftliche Expedition nach den nordwest- lichen Provinzen Chinas, Insecten, 41. Hymeno- ptera, 6. Sphegidae. — Arkiv for Zoologi (27A (21): 1—15, figs. 1—6. Iwata, K., 1939. Habits of some solitary wasps in For- mosa. IV. — Transactions of the Natural History Society of Formosa 29: 161—178, 3 figs. Kohl, F. F., 1918. Die Hautflüglergruppe Sphecinae IV. Die natürliche Gattung Sceliphron Klug (Pelo- poeus Latr.). — Annalen des Naturhistorischen Museums in Wien 32: 1—171, figs. 1—82. Lepeletier de Saint Fargeau, A., 1845. Histoire natu- relle des insectes, Hymenopteres 3: 1—644. — Roret, Paris. Maindron, M., 1878. Notes pour servir à l’histoire des Hyménoptères de l’Archipel Indien et de la Nou- velle-Guinée. I. Observations sur quelques Sphegiens (G. Pelopaeus) de l’Archipel Indien. — Annales de la Société Entomologique de France (5)8: 385398, pl. 9. Michener, C. D., 1944. Comparative external mor- phology, phylogeny and a reclassification of the bees (Hymenoptera). — Bulletin of the American Museum of Natural History 82: 151—326, figs. 1—246, diagrams 1—13. Mocsary, A., 1883. Hymenoptera nova europaea et exotica. — Ertekezések a Természettudomanyok Köreböl 13: 1—72. Morawitz, F., 1888. Hymenoptera aculeata nova. — Trudy Russkago Entomologicheskago Ob- shchestva 22: 271. Rohwer, S. A., 1921. The Philippine wasps of the sub- family Sphecinae. — Philippine Journal of Science 19: 665—676. 260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 8, 1986 (1987) Smith, F., 1856. Catalogue of the Hymenopterous in- sects in the collection of the British Museum, iv, Sphecidae, Larridae and Crabronidae: 207—497. — London. Smith, F., 1858. Catalogue of the Hymenopterous in- sects collected at Sarawak, Borneo; Mt. Ophir, Malacca; and at Singapore, by A. R. Wallace. — Journal of the Proceedings of the Linnean Society of London, Zoology 2: 42—130. Smith, F., 1859. Catalogue of the Hymenopterous in- sects collected at Celebes by Mr. A. R. Wallace. — Journal of the Proceedings of the Linnean So- ciety of London, Zoology 3: 4—27. Smith, F., 1861. Descriptions of new species of Hy- menopterous Insects collected by Mr. A. R. Wal- lace in the Islands of Batchian, Kaisaa, Amboyna, Gilolo and at Dory in New Guinea. — Journal of the Proceedings of the Linnean Society of Lon- don, Zoology 5: 93—143, pl. 1. Smith, F., 1863. Catalogue of the Hymenopterous In- sects collected by Mr. A. R. Wallace in the Islands of Mysol, Ceram, Waigiou, Bouru and Timor. — Journal of the Proceedings of the Linnean Society of London, Zoology 7: 6—48. Smith, F., 1870. Descriptions of some new species of Apidae and Vespidae collected by Mr. Horn. — Transactions of the Zoological Society of London 7: 161—196, pls. 19—22. Strand, E., 1914. Ueber das Nest von Sceliphron de- forme Sm. — Archiv für Naturgeschichte 80A (10): 116—117. Strand, E., 1915. Ueber einige orientalische und pal- aarktische Crabroniden der Gattungen Sphex, Sce- liphron und Ammophila im Deutschen Entomolo- gischen Museum. — Archiv fur Naturgeschichte 81A(5): 88—97. Tsuneki, K., 1964. Some wasps collected by Dr T. Shirözu in Kyushu. — The Life Study, Fukui 8: 10. Tsuneki, K., 1971. Studies on the Formosan Sphecidae (IX). A supplement to the subfamily Sphecinae (Hym.). — Etizenia 53: 1—7, figs. 1—14. Tsuneki, K., 1972. On some species of the Japanese Sphecidae (Hym.), notes and descriptions. — Eti- zenia 59: 1—20. Tsuneki, K., & K. Shimoyama, 1963. Sphecidae and Chrysididae collected in the district of Towada, | Japan. — The Life Study, Fukui 12: 48—50. | Turner, R. E., 1912. Notes on fossorial Hymenoptera, | VI, on the species collected in New Guinea by the | expedition of the British Ornithologists Union. — | Annals and Magazine of Natural History (8)9: 194—202. Turner, R. E., 1917. On a collection of Sphecoidea | sent by the Agricultural Research Institute, Pusa, Bihar. — Memoirs of the Department of Agricul- | ture in India, Entomological Series 5: 173—205. Uchida, Y., 1925. A list of known species of the Co- rean Hymenoptera which I collected in 1922, and their geographical distribution. — Insect World | 29: 328—337, 366—373. | Vecht, J. van der, 1957. The Sphecoidea of the Lesser Sunda Islands (Hym.) I, Sphecinae. — Verhand- lungen der Naturforschenden Gesellschaft in Basel 68: 358—372. Vecht, J. van der, 1961. Ueber Taxonomie und Evolu- tion der Grabwespengattung Sceliphron Klug. — Verhandlungen XI. Internationaler Kongress für Entomologie, Wien 1: 251—256, figs. 1, 2. Vecht, J. van der, 1984. Die Orientalische Mauer- wespe Sceliphron curvatum (Smith, 1870) in der Steiermark, Osterreich. — Entomofauna 5: 213— 219, figs. 1—5. Vecht, J. van der, & F. M. A. van Breugel, 1968. Revi- sion of the nominate subgenus Sceliphron Latreille (recte: Klug) (Hymenoptera, Sphecidae) (Studies on the Sceliphronini, part I). — Tijdschrift voor Entomologie 111: 185—255, figs. 1—27. Wiley, E. O., 1981. Phylogenetics, the theory and practice of phylogenetic systematics: xv + 1— 439. — New York, etc. Williams, F. X., 1919. Philippine Wasp Studies. — Proceedings, Hawaiian Sugar Planters’ Associa- tion Experiment Station, Entomology 14: 1—186. Yasumatsu, K., 1942. Hymenoptera aculeata collected by Mr. K. Tsuneki in North China and inner Mongoiia. I. Sphecoidea. 1. List of species. —Mu- shi 14: 103—104. R. V. Hensen: The subgenus Prosceliphron 261 INDEX Be iE Maindron).........eeeesssacserscennnreeene. 254 Le (MO EM) ene 240 Prema ae an der Vecht 2... 249 MENTON)... 256 Hoyt CSM 256 coromandelicum (Lepeletier)....................... 243 EOS RER RES 228 He (Sid inn 230, 236 aller Ko En 256 Aeon SONORE 236 rans SIN) iii 242 [izuofasciaium: (Smith)... 249 Nonas (Smith) pini 255, 258 Pret tone inn 226 UOMO nn 240 RAS D oem BS) Ds Ale: dns 249 RorcanuingUchiida nme. 240 Tatieanetue SSPAn Renee herren 252 Voen tine ne am enon re nn 234 moes AD al MOGEN: 254 MU Tann Smith) E nn 254 PIREO n 240 ocelli ene 258 JODO LID CES NRE 258 palchi Gissi 228 AED) nl ELSE ER 228 nes (Mocsany) ner 254 open Sichere 247, 249 CHER ODA CUS AROMNS Kill teeta ee 224 ALTO ans Ken: 238 HD Ale CAME ONE een 234 HI AS CLALIT: SM 254 ORTEN 3 eV NE nn. SR Da .z sa è oa t N ial a ae Sime T B ud ae Ni La : 6 Le ara Wao nr le; LEA me no al Tiss Si lid | Ra DEEL 129 AFLEVERING 9 1986 AL UGI me TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING INHOUD M. A. Liertinck. — New and little known Platycnemididae and Coenagrionidae from New Guinea and the Solomon Islands (Odonata), pp. 263—291, figs. 1— 43. Tijdschrift voor Entomologie, deel 129, afl. 9 Gepubliceerd 31-VII-1987 NEW AND LITTLE KNOWN PLATYCNEMIDIDAE AND COENAGRIONIDAE FROM NEW GUINEA AND THE SOLOMON ISLANDS (ODONATA) PF mg VIE by ( SMITHSON. OAL / 14 LPS VAS ; VATA M.A. LIEFTINCK + \ N . > è, y 7 With notes added by J. van Tol 7 + Rijksmuseum van Natuurlyke Historie, Leiden, The Netherlands RIES ABSTRACT Five new species of Platycnemididae (Odonata), viz., Rhyacocnemis prothoracica, Salo- mocnemis gen. nov. gerdae, Lieftinckia malaitae, L. isabellae and L. ramosa, and four new species of the genus Teinobasis (Coenagrionidae), viz., T. simulans, T. obtusilingua, T. chionopleura and T. imitans, are described. Papuagrion gurney: Liettinck and Teinoba- sis emarginata Lieftinck are synonymized with Teinobasıs aluensis Campion. Besides sever- al other species of these families from the Solomon islands are discussed. This posthumous publication concludes the scientific odonatological studies of Dr M. A. Lieftinck. [INTRODUCTION] by J. van Tol After the death of Maurits Anne Lieftinck on April 12th, 1985, most of his scientific material was donated to the Rijksmuseum van Natuur- lijke Historie (Leiden). I have thought it one of my first tasks to investigate whether the manu- scripts that Dr Lieftinck had at hand, could be prepared for publication. During my last visit to him in February, 1985, he showed me drawings and manuscripts in preparation. One large manuscript on the genus Procordulia and allied genera, for which many drawings were already prepared, is not available for publication; the second manuscript has resulted in the present paper. The manuscript appeared to be almost com- plete, and thus suitable for publication. The drawings and photographs were all mounted on cardboard, and numbered. The descriptions of all but one species were available, but a general introduction, an introduction to the Platycne- mididae and concluding remarks were lacking or not in a form ready for publicaton. I have added the minimum amount of text, restricting myself to additions that were needed to publish a consistent paper, viz., a title, an abstract, an introduction, a list of species included, a few ti- tles of chapters (i.e. names of genera), the refer- 263 ences in the next to the figures, the captions of the figures, and the references. No serious problems were met with this work. The number of scientific papers in this field is rather limited, so that I could easily find the works referred to. The captions to the figures were prepared from notes made in pencil with the drawings, as well as from the text. Since the description of the specimens from the Thomas W. Donnelly col- lection, numbered “72 x 040”, was lacking, I have left out the preliminary new genus and species name for this taxon, which is closely re- lated to Torrenticnemis. The figures, which were already mounted by Dr Lieftinck, are, however, included. The names of chapters added by me are indicated in square brackets. I should like to emphasize explicitly here, that Dr Lieftinck should be considered the au- thor of all new names introduced in this paper. [| ACKNOWLEDGEMENTS | The cooperation of the curators of the Odo- nata departments of the following institutions is gratefully acknowledged (the abbreviations used for the collections are given in brackets): Bernice P. Bishop Museum, Honolulu (BISH), British Museum (Natural History), London (BM), Muséum National d’Histoire Naturelle, Paris (MP) and United States National Museum, Washington (USNM). 264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) Soon SSS = Sif Lie [> ON ET Figs. 1-4. Wings of Platycnemididae. — 1, Idiocnemis obliterata Lieftinck, 1932; 2, Paramecocnemis erythro- stigma Lieftinck, 1932; 3, Lochmaeocnemis malacodora Lieftinck, 1949; 4, Torrenticnemis filicornis Lieftinck, 1949. LIEFTINCK: New Guinea and Solomon Islands Odonata 265 The Rijksmuseum van Natuurlijke Historie, Leiden, is abbreviated as ML. Special thanks are due to Dr Thomas W. Donnelly (Binghamton, N.Y.) for making spec- imens from his own collection available for study. [LIST OF SPECIES INCLUDED] Platycnemididae Genus Rhyacocnemis Lieftinck, 1956 R. prothoracica spec. nov. Salomocnemis gen. nov. S. gerdae spec. nov. Genus Lieftinckia Kimmins, 1957 L. salomonis Kimmins, 1957 L. lairdi Lieftinck, 1963 L. malaitae spec. nov. L. kimminsi Lieftinck, 1963 L. isabellae spec. nov. L. ramosa spec. nov. Lieftinckia spec. indet. 1 Lieftinckia spec. indet. 2 Genus Coenagrionidae Teinobasis Kirby, 1890 T. aluensis Campion, 1924 T. simulans spec. nov. T. obtusilingua spec. nov. T. chionopleura spec. nov. T. imitans spec. nov. T. bradleyi Kimmins, 1957 Genus PLATYCNEMIDIDAE Rhyacocnemis Lieftinck, 1956 Rhyacocnemis prothoracica spec. nov. (figs. 5, 8) Material. — NE.New Guinea: 1 d (adult, holo- type), Morobe Dist., stream at Gurakor, Wau Rd., 500 m, 25.x.1972, nr. 72X044, Thomas W. Donnelly leg., in coll. ML. Male (adult, holotype). — Smaller than R. sufficiens Lieftinck, 1956. Head short and broad, width across eyes only 4.0 mm. Labium bright creamy yellow, median lobe with short V-shaped emargination about 1/5 as deep as midlength of lobe itself, its apical lobes some- what convex exteriorly, obtusely rounded off, tips, as also the outer border of palpi, fringed with ferruginous bristles. Labrum and clypeus broad and prominent as in sufficiens, well visible from above; labrum convex and widest ante- riorly, its surface smooth and rather shiny, deep black; anteclypeus pale at middle, postclypeus with pair of bright yellow transverse spots, one each side, which taper inward leaving a thick T- shaped black mark occupying most of the sur- face. Frons in front and whole genal area taken up by a sharply defined chrome yellow bar con- necting the eyes, which extends upward to al- most reaching the antennal sockets; remainder of head above and underneath deep coal back with no other pale marks or pruinescent areas than a pair of large, isolated, pear-shaped sky- blue postocular spots pointing inward. Anten- nae long (about 2.5 mm), black; scape cylindri- cal, short and thick, only little longer than its diameter at apex, pedicel slender, slightly clubbed, about 7—8 times as long as its apical width, third segment much thinner but of equal length, the distalia long and thread-like. Prothorax markedly dissimilar to that of R. sufficiens; anterior lobe long, rather de- pressed and lacking a distinctly swollen anterior rim, its dark surface indistinctly mottled with ferruginous specks and with a pale margin; pro- notal tubercles strongly raised, forming a pair of robust slightly diverging cone-shaped and bluntly pointed processes whose surface is dull and coarsely wrinkled; these pyramidal bosses are about equal in height to the smooth pro- pleurae and directed obliquely upward and backward; posteriorly, they are feebly ridged on either side, enclosing a more flattened for- ward slanting posterior face; immediately be- hind each of them arises a small and thin trans- verse plate, situated on a much lower level; pos- terior lobe broad and collar-shaped, about equal at midlength to the anterior lobe by lying flatly down, its side angles triangular, slightly project- ing and a little downbent. Colour of whole dor- sal surface of prothorax lustreless dark brown on coarsely wrinkled ground, all lateral parts re- maining smooth and contrastingly coloured a bright greenish yellow. Mesothorax with a complete, almost parallel- sided, dull bronze-black middorsal band occu- pying the inner halves of mesepisterna, the mid- dorsal carina and ante-alar triangles being also black; outer halves of mesepisterna lighter, forming a dark ferruginous brown juxtahumeral band equal in width to the black middorsal one, this coloured area acquiring a much lighter tint upon lower 1/4 of mesepimera in the form of an 266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) 3 Figs. 5—6. Male appendages of Platycnemididae in dorsal and left lateral view. — 5, Rhyacocnemis prothoracica spec. nov.; 6, Unnamed species, “72040” collection of Donnelly. elongate patch of bright chrome which is con- tinued ventrad upon the mesostigmal laminae. A black line, broadening at lower end to occupy most of the mesinfraepisternites as well, accen- tuates the humeral suture. Sides, as far down and just including the metaspiracle, bright chro- me intermingled with light green; behind this, an almost equally broad but very irregular band of rusty dark brown runs over the second su- ture, expanding ventrad to cover also most of the metinfraepisternites; rest of metepimeron distinctly light blue, the lateroventral carina and whole vental surface of thorax, pale yellow; poststernal plate almost bare, but provided upon its middle with a compact patch of many longish, dark, stiff bristles placed upon a dis- tinctly swollen part of the sclerite, the setae making up this paintbrush-like tuft being di- rected straight back though remaining well visi- ble in profile view. Legs thin and slender, hind femur reaching back to a little beyond halfway length of first abdominal segment; femoral bristles 11—12 each side, longest on hinder pair, almost three times the interspaces but decreasing in length toward base, 8—9 on mid and fore femora; tib- ial bristles longer, 7—8 in outer row of hinder pair; tarsi slender, the claws with small but acute subapical inferior tooth. Colour of all coxae and trochanters blue-green, outer faces of all femora with a thick black stripe broadening gradually toward end including the knees, inner faces bright chrome darkening toward apex; all tibiae and tarsi black but claws yellowish. Wings (fig. 8), narrower with more pointed tips than in both /diocnemis and Paramecocne- mis (figs. 1, 2), shape and venation very similar to R. sufficiens (fig. 7), the undulations of post- pterostigmal border slightly more pronounced than in the latter but less so than in /diocnemis; neuration black. Pterostigma oblique, a trifle longer than high, dark brown heavily framed in black. Wing bases with all axillary sclerites as well as the meso- and metanota strikingly col- oured a brilliant sky-blue. Abdomen (end segments fractured), very slim and slender but not unusually drawn out and lacking conspicuous hair tufts on any of the ter- gites such as seen in both species of male Para- mecocnemis. Structural peculiarities are (1), postero-lateral edge of tergite 2 produced back- ward, ending in a short triangular lobe (instead of normally cylindrical and parallel-sided in suf- ficiens), its lower border sparsely beset with golden yellow setae; (2), Antero-lateral edge of tergite 3 with a short, nipple-shaped protuber- 267 LIEFTINCK: New Guinea and Solomon Islands Odonata (ima >| See, u Ener] Figs. 7—9. Wings of Platycnemididae. — 7, Rhyacocnemis sufficiens Lieftinck, 1956; 8, Rhyacocnemis protho- racica spec. nov.; 9, Unnamed species, “72 X40” in Donnelly collection. 268 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) ance directed laterad (this tergite normal in suf- ficiens); intermediate segments thin and needle- like, but from end of 7 on gradually expanding in both dimensions, 8 about one and one-third times longer than 9, the latter squarish and par- allel-sided, 10 short and broad. Almost wholly black; tergites 1 and 2 conspicuously marked with sky-blue and bright ochreous: distal half of 1 blue, finely interrupted by black in the median line, this stalk attached to a narrow black apical ring; 2 on either side with complete, thick, bright chrome lateral bar occupying lower por- tion of tergite; next segments, inclusive of the anal appendages, apparently all black (tergites 8 and 9 possibly discoloured and partly blue in life?), save merest traces of obscurely coloured basal annules and a yellow line bordering ven- tral margin of tergites 3—7. Directly from be- low mid-apical border of tergite 10 projects a slender, rod-like cylindrical process whose apex is subtruncated and a little incised. Anal appendages as in fig. 5; superior pair about equal in length to the subquadrangular 9th segment, at first outcurved and bluntly toothed on the inside near base, then broaden- ing, bent inward and somewhat flattened, strongly transversely ridged on uneven ground, inner border in dorsal view irregularly sculp- tured and toothed, outer faces of basal portion of each beset with short and strong spike-like denticles interspersed with longish hairs (only partly shown in dorsal view and omitted inside view sketch). Inferior appendages well devel- oped but attaining only half the length of supe- rior pair, basal portion of each broad and rather swollen, then rather suddenly narrowing and somewhat curved toward each other, gently ta- pering to slightly upcurved tips. Salomocnemis gen. nov. Stature of Lieftinckia, but with a broader head and more bulging eyes. Labium similar, the median emargination equally small but less narrow, the limbs of the U distinctly diverging instead of subparallel. Shape of mouth-parts and face much as in Lieftinckia salomonis (the type- species), but all parts comparatively a little broader. Anteclypeus forming a right angle with the postclypeus, the latter distinctly carinate an- teriorly. Frons obtuse-angulate anteriorly with well-pronounced though rounded, transverse ridge, instead of gently sloping down and rather flattened in Lieftinckia; frontoclypeal suture broad, impressed, rectangulate, the sulcus be- tween antennal sockets likewise distinctly im- | pressed. Inner orbital line straight (not convex), strongly converging anterad on dorsal as well as on ventral surface of head (parallel-sided ven- | trally in Lieftinckia). Antennal sockets but little raised but distinctly swollen; scape only slightly longer than its socket, thick, cylindrical, little | more than twice as long as broad; pedicel more slender and rather clubbed apically, subequal in length to scape; segment 3 thinner and about | one-fourth shorter than pedicel, remaining seg- | ments about equal in length to scape and pedicel united. Vertex small, raised, ocelli in equilateral | triangle. Occipital ridge subacute. | Prothorax not modified, pronotal tubercles | raised, conical; posterior lobe broad, surface | convex, rather dome-shaped, side-angles round- | ed off in both sexes. Legs much shorter and less | slender than in Lieftinckia; hind femur not | nearly attaining apex of 2nd abdominal seg- | ment; armature as in the allied genus, but all | femoral and tibial bristles comparatively a little shorter; tarsal claws with distinct subapical in- ferior tooth only little shorter than the claw it- self. Wings less abruptly petiolated, more evenly broadened and with more open neuration than in Lieftinckia. Nodus situated more distad, ap- proximately one-third of the distance from base to apex. Arculus at Ax,; Ac placed distal to Ax, instead of far proximal to that level in Lief- tinckia; petiolation of the wing as in that genus ending at a point where Ab meets the wind bor- der, but Ab and Ac coincident at margin instead of far apart, Ab occasionally terminating slight- ly proximal to Ac, almost at level of Ax,. Rs at or slightly distal to subnodus, M, well before that level, the distance separating them at origin shorter in fore wings than in hind wings. Two postquadrangular antenodal cells. Quadrilateral hardly broadened distally, costal side much lon- ger than distal side, especially in hind wing. Me- dio-anal link entire, or almost so. Wing border of whole apex posterior to pterostigma, back to termination of Cu,, slightly but distinctly undu- lated (most marked on hind wing). A single row of cells between C and R, beyond pterostigma. Pterostigma rather swollen, almost square, cov- ering one underlying cell. Abdomen slender, basal and terminal seg- ments moderately inflated. Proportionate lengths of segments as in Lieftinckia. Colour pattern much as in that genus. Vesica spermalis of d penile organ not prominent, suboval, thin and plate-shaped in profile view, membranous LIEFTINGK: New Guinea and Solomon Islands Odonata 269 Fig. 10. Wings of Salomocnemis gerdae spec. nov. (Guadalcanal). in the centre, but with strongly sclerotised lateral rims; ligula (penis) shaped as in fig. 15; no shaft spines. Male anal appendages of rather simple structure, resembling L. salomonis. Eighth abdominal sternite of female simple, without any indication of an apical spine; geni- tal organs normal, valves surpassing tenth seg- ment and tuberculum anale for about the same length as the former, lower margin of outer valves finely denticulate; cerci short and blunt, flattened dorsoventrally. Type-species: Salomocnemis gerdae spec. nov. Habitat: Solomon Is. (Guadalcanal). Salomocnemis gerdae spec. nov. (figs. 10, 14—16) Material. — Solomon Is.: 2 & 1 © (1 d subadult), one pair of juveniles, Guadalcanal I., Komugelea, 1200 ft., “in forest”, 22.1x.1965, R. Slooff and Gerda Stanny Slooff-de Vries (all ML). Male (subadult, holotype). — Head with labi- um palest yellow, mentum and centre of median lobe semitransparent, colourless. Mandible- bases dark yellow-green, the teeth black. La- brum, clypeus, genal area and frons, dark olive- green, rather shiny; all the rest of dorsal surface of head, including the antennae, similarly col- oured but dull, sparsely marked with light brown, as follows: a minute central spot in the depth of ante-ocellar depression, basal half of antennal pedicel, a crescentic spot to the inside of each lateral ocellus, and a vestigial spot upon middle of occipital ridge; a curved black line ex- tends inward on either side of middle upper or- bital margin as far as outer end of occipital ridge, the upper part of the postocular area be- hind this line being olive-green, the lower por- tion dark brown fading to dirty yellowish un- derneath. Prothorax, inclusive of anterior and posterior lobes, dull orange, growing paler and inter- mingled with light green laterally; pronotal tu- bercles little elevated, rounded; posterior lobe depressed, at least four times broader than long, surface slightly convex and undulated, its poste- rior border a little upturned, almost straight, side-angles rounded. Synthorax orange, deepest in tint at the humeral suture but becoming light- er with admixture of pale green upon lower portions of epimera and infraepisterna; dorsum, to a little in advance of humeral suture, olive- green; mesoprescuta orange, with deep black speck on either side of the triangles, the latter also orange but with their borders deep black, as are the postero-dorsal edges of the metepime- ron. Legs with coxae and trochanters palest orange, for the rest light greenish to greyish yel- low; apices of all femora and last tarsal seg- ments, narrowly ringed with brown; all bristles and teeth of tarsal claws brown. Wings hyaline, the open neuration black, on- ly the petiole somewhat more palely veined. Whole margin of wing apex posterior to ptero- stigma with a number of small but distinct un- dulations, the posterior border of hind wing 270 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) moreover noticeably protruding at a point where M, and Cu, enter the margin. Fore wing with 14, hind wings with 12 Px of first series; Rs arises very slightly distal to subnodus, M, well proximal to it; M, at Px, in fore wings, a little beyond Px, in hind pair; M,, three and four cells further distad, respectively. (In all fore wings and in four out of six hinds wings, the anal bridge terminates even a little proximal to Ac, which in the female is placed under Ax, instead of slightly distal to it). Abdomen, ground colour yellowish, distal portions of tergites 3—7 marked indefinitely with brown, this colour progressively becoming more extensive and darker posteriorly; hind margins of 1—2 finely ringed with black and each with a dark middorsal line; basal 1/3 to 2/5 of 3—7 yellow, passing gradually to brown pos- teriorly, the apical rings deep black; 8 black, 9 brownish black mottled with yellow, dorsum of 10 entirely yellow, the sides greenish with a deep black streak alongside middle of posterior border. Anal appendages (fig. 14), superior pair trian- gular, acutely pointed, inner faces of each rather flattened, the basal portions also triangular, di- rected obliquely mesad and ventrad, tapering to a blunt black-tipped process visible only in cau- dal view; inferior appendages also yellowish, rather longer, strongly hollowed out within, the apices of each incurved, squarely cut off and furnished with a row of five black denticles. Female (juvenile). — N.B. Where not men- tioned in the description of the male, most of the next details of structure also apply to that sex. Resembles the male in most respects. Mouth- parts in better condition than in the available males. U-shaped median emargination of labi- um elongate, about 1/5 total length of midlobe, the U itself parallel-sided; colour of whole structure as well as mandible-bases, palest chro- me. Labrum porrect, slightly more than 1% times broader than long, widest at middle, its surface convex, smooth. Structure of all upper parts of head and antennae as described for the male, coloration a pale creamy yellow inclusive of the sockets and scape of antenna, the pedicel and third joint being grey-black, for the rest dis- talia again lighter. Postocular area cream, except its anterior suture finely deep black; occipital region gradually changing to brown. Prothorax much as in male: surface smooth, pale ochreous almost throughout; anterior lobe broad, of the usual shape, subequal in width to the posterior one but strongly swollen in side view, moderately raised, its front border thick- ened, straight-lined and subacute in dorsal view, but protuberant in side view, with pair of deep black transverse rims one each side; pronotum with three pairs of somewhat convex, roundish tubercles placed in the long axis, the outer pair largest, the innermost pair diverging anterad but fused together in the median line posteriorly, the outer tubercles covered with extremely min- ute, finely pointed tubercles; transverse sulcus separating them from posterior lobe moderately deep, surface irregular. Posterior lobe of pro- thorax quite simple, not at all modified, mid portion somewhat raised, bow-shaped in caudal view, the whole structure about 4—5 times broader than long, hind margin subacute, al- most straight at middle in dorsal view, sides gently rounded, hardly downcurved. Mesostigmal laminae small, subtriangular, yellow, each with a raised, deep black inner rim placed in the long axis, lateral parts triangularly tapered (fig. 16). All colour marks on dorsum of synthorax ill-defined, pale chrome with faint greenish hue: a pair of wedge-shaped antehu- meral bands, widest below, tapering and bluntly pointed upward, on a pale ferruginous ground, this colour extending laterad to almost halfway metaspiracle; ante-alar triangles lighter but fine- ly black-rimmed, as are also the dorsal ridges of mesopleurae, a speck at upper end of second su- ture, and the posterior borders of the metepime- ron. Legs throughout light greenish yellow, the femoral banding hardly indicated; all spines and bristles obscured; tarsal claws as described for the male: with small but distinct, sharply point- ed dark inferior tooth very near the apex prop- ete Wing neuration light brown, membrane hya- line. Venation, inclusive of the relative positions of Ac and origin of M; — Rs as well as shape of pterostigma almost exactly as described for the male (fig. 10), but differing from the latter in that the wing tips are more bluntly rounded, the post-pterostigmal as well as the underlying cells being markedly higher, those of the latter being subquadrangular or even higher than long (instead of the reverse in two males). Also, the marginal undulations at wing tips and end of main veins, are distinctly more pronounced than in the opposite sex. Pterostigma dark grey, con- trastingly framed in bright yellow. Abdomen slender, of the usual shape, basal segments scarcely broadened, thereafter grad- LIEFTINCK: New Guinea and Solomon Islands Odonata 271 ually a little expanded toward the end (imma- ture specimen!). Ground colour cream, marked indistinctly with brown: traces of that colour on middorsum and sides of tergite 2, whole 4/5 of 3, 2/5 of 4—6, almost distal half of 7, and most of 8 including the valves; 9—10 as well as the cerci and tuberculum, all yellow. Genital valves | rather long, projecting beyond apex of abdo- men, equal in length to lower margin of tergite 8; ventral border of valves only slightly convex in side view, apical 3/5 of each armed with a row of ca. 24—26 microscopical swollen tuber- cles gradually increasing in size toward apex, each of the most distal ones ending suddenly in a minute acuminate spine. Measurements: ¢ abdomen + appendages 34.6 mm, hind wing 21.5 mm; 2 abdomen (incl. valves) 33.1 mm, hind wing 21.4 mm. Lieftinckia Kimmins, 1957 Lieftinckia salomonis Kimmins (fig. 17) Lieftinckia salomonis Kimmins, 1957: 312—315, fig. 1 A—G (89 struct.), &? Guadalcanal. — Lief- tinck, 1963: 527—530, 541—542 (8? incl. key), 531—533 (6 larval struct.), pl. 26 fig. 1 (d wings), tfig. 1—12 (6 imaginal char. & larval struct.), 3 2 Guadalcanal. Additional material. — Solomon Is.: 1 2 (juvenile) Guadalcanal I., Tenaligi River 25.ix.1965, “heavy for- est”, R. Slooff (ML); 1 & (adult), Guadalcanal I., Gal- lego, 11.vii.1965, “hidden valley, 2nd grass clearing”, Roy. Soc. Exped. Brit.Mus.1966 (BM); 1 d 2 2 (sub- adult), Guadalcanal, Gold Ridge, 22.11.1955 (4) and Tsarivonga river, same date (©), E. S. Brown (BM). Male. — The Gallego specimen from Guadal- canal is the first full-coloured male of the type- species of Lieftinckia. Like so many other platy- cnemidids, salomonis also passes through a se- ries of colour phases before reaching maturity, the present case being a good example of this extraordinary variation. With its pale brown and cream-coloured body, the present male looks quite different from the immature insect. Labium obscurely red-brown, all the rest of the face vividly brick red as far upward as a little behind the ocellar triangle; thereafter the red changes abruptly to black in a line that runs from eye to eye, passing over the occipital crest, which itself is dark brown in the middle. Hind portion of the postocular lobes and rear of the head, deep black except a triangular spot on the lower genal area. Dorsum of prothorax and up- per part of its sides dull black, for the rest obscurely orange-red. Mesepisternites and ante- alar triangles black, almost lustreless, this colour passing beyond the humeral line for a short dis- tance parallel to the suture but suddenly ex- pands upward so as to occupy also the dorsal one-third of the mesepimeron. Sides and ventral surface of thorax bright chrome, only most of the infraepisternites rusty brown. Coxae and femora inclusive of the bristles, orange-red; outer faces of fore femora more obscured, knees also darkened; tibiae and tarsi reddish black. Wing membrane suffused with grey-brown, pterostigma brownish black. Abdomen dark brown becoming black on the terminal seg- ments; sides of 1 and those of 2 for the basal 2/3 part, ochreous; fairly distinct basal and subter- minal spots of the same colour bordering the tergites laterally being also present on 3—6, these spots elongate and fused together on 7. The three further Guadalcanal specimens (Ta- penanje), are near topotypes collected in 1955. These were found unnamed in the British Mu- seum collection years later than the examples of either sex discussed earlier by Kimmins (1957) and myself (1963). I made the following brief colour note on the subadult male: “Head bright orange, dorsum of synthorax black, the sides and legs orange; ab- domen obscured, the annules sharply defined, segm. 10 and anal appendages yellow”. Lieftinckia lairdi Lieftinck Lieftinckia lairdi Lieftinck, 1963: 534—537, 542 (key d 2), pl. 26 fig. (d wings), tfig. 13—16 (d struct.), 3 2 Guadalcanal. Lieftinckia spec.? Kimmins, 1957: 315, 2 Guadalca- nal. Additional material. — Solomon Is.: 1 3 (adult), Guadalcanal I., Sutakiki River, 23.11.1955, E. S. Brown, no. 2605 BM 1957—201 (BM); 1 & (sub- adult), Guadacanal I., Nuhu, 26.x.1965, “slow flow- ing pool formed by seepage alongside river, by ford”, Roy. Soc. Exped. Brit. Mus. 1966 (ex BM, ML). Male (adult, Sutakiki river). — This is a near topotype, larger in size than the previously de- scribed immature specimens. The following de- tails are noteworthy additions to the original description. 272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) Prothorax shaped as in the immature insect but colours generally darker; posterior lobe more evenly bordered, showing no undulations such as seen in the juvenile males. Sides of syn- thorax red-brown, the meso- and metepimera predominantly light green. The subapical dark rings at the femora are broad but not sharply pronounced. Wings relatively shortly stalked, neuration likewise not differing from that origi- nally described and photographed (loc. cit., 1963, pl. 26 fig. 2). Fore wing with 22—23, hin- der pair with 20 Px of first series; M, arises just before the subnodus, Rs half a cell beyond that level. The short, narrow and very oblique pte- rostigma approaches salomonis most closely in shape. Abdominal tergites mainly blackish brown, but dorsal bands much less constricted and con- trastive with the light ground colour, than in malaitae spec. nov., described hereafter. Measurements: abdomen + appendages 40.5 mm, hind wing 28.0 mm. So far only known from Guadalcanal. Lieftinckia malaitae spec. nov. (fig. 12) Structurally very similar to L. lairdi, but ob- viously specifically distinct from that species. Material. — Solomon Is.: 2 & 1 2 (semiadult), Ma- laita I., Dala, 50 m, 9—14.vi.1964, J. & M. Sedlacek (Gz WSs, MIDs De @ ack, i euvel), Melk Ie Ngwaiau, 1500 ft., 10.x.1967, R. Slooff (holotype d and 2 paratypes, ML). Male (adult, Ngwaiau). — Antenna long, al- most equal in length to inner orbital line when viewed from above; scape short and cylindrical, little longer than broad, pedicel thin and much slenderer (apparently a little varying in length: in one paratype fully three times longer than scape), both segments pale in colour; distalia darkened, about as long as first two segments united. Head, thoracic sclerites and legs, all more uniformly coloured than in lairdi, of a delicate cinnamon to light brownish orange, deepest in shade on top of frons, the ocellar region and on the thorax above; otherwise unmarked save minute dark lines and specks at the median cari- na, dorsal margin of mesepimeron, the upper- most edges of the pleural parts, and at the lateroventral angle of metepimeron. Whole ven- tral surface of thorax as well as the coxae, tro- chanters and femora, throughout pale, the ex- treme tips and much broader subapical rings of | femora brown, the latter distinct though not sharply outlined; tibiae, tarsi and bristles slight- | ly more obscured than the femora. Shape and neuration of wings as shown in fig. 12, relatively narrow and elongated, more | abruptly stalked than in salomonis and lairdi, | the stalk also being longer than in the latter, | while the pterostigma is slightly less oblique, higher and more nearly parallel-sided. Fore wing with 18—22, hind wing with 17—18 Px of first series. Ground colour of abdomen light ochreous, but tergites 2—7 much more contrastingly marked with blackish brown than in any other species. Dorsal bands extending from base to apex, forming complete, fairly distinct marks, which toward the base of each tergite are rather pointed, leaving elongate yellow lateral spots, but distad are markedly constricted before be- coming fused with well-defined black apical rings. Tergites 8—10 much obscured, only the sides usually broadly yellowish. Anal appendages (not figured) yellow, the su- periors darkened above, shaped much as in Jair- di, but differing as follows: superior appendages triangular, rather inflated basally, as in laırdi, but more distinctly pointed apically; inferiors less strongly upcurved with the black tips sharply hooked inward and acuminate, instead of tapering gradually, as seen in lairdi. The ro- bust interobasal spurs of the superior append- ages are distinctly longer than in lairdi and wholly exposed in side view, at first directed straight ventrad, then gently curving cephalad; in caudal view these processes are directed to- ward one another, the apices meeting at their extreme tips, thus remaining uncrossed. Female (subadult). — Resembles the male in most respects but differs in details of colouring and the better defined abdominal pattern. Mouth-parts pale yellow, the labrum mixed with light ochreous; face, the swollen antennal sockets, and an ill-defined broad area along in- ner orbits tending to palest green, the vertex surrounding the ocellar area and most of the oc- cipital region behind it, dark cinnamon, this col- our occupying most of the dorsal surface be- tween the eyes posterior to the lateral ocelli. Antennae pale, shaped as in male, the long slen- der pedicel tipped with black. Ventral surface of head palest greenish yellow. Pro- and synthorax light brownish yellow, rather darkened upon all upper parts: brown LIEFTINCK: New Guinea and Solomon Islands Odonata 273 Figs. 11—12. Wings of Lieftinckia species (Platycnemididae). — 11, L. isabellae spec. nov. (Santa Isabel); 12, L. malaitae spec. nov. (Malaita). becoming dirty orangish on the surtace on ei- ther side of the humeral line, and still more dif- fusely so along the second suture, between these darkish areas is a band of pale yellowish green. Prothorax little modified, pronotal tubercles low, not at all prominent, their dorsal line in side view almost straight; anterior lobe raised, of the usual shape, its border simply convex; posterior lobe short and broad, shorter than the pronotum itself, slightly raised and almost straight-lined in dorsal aspect but hind border distinctly undulated with obtuse-angulate dor- solateral edges, the little prominent lateral lobes small, narrowly rounded off. Legs including bristles pale, femora ringed as in male, brownish. Wings hyaline, the apical crenulations as dis- tinct as in male. Fore wing with 19, hind wings with 17 Px of first series. Pterostigma shaped exactly as in the opposite sex, i.e. with sides more nearly parallel than in Jairdi; colour pale grey-brown. Abdomen slender but less so than in male, with cylindrical segments of almost even width, except the last ones very slightly expanded. Ground colour light ochreous, the dorso-lateral dark brown marks much more sharply defined, standing out more clearly than in male, as fol- 274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) lows: tergite 1 only the raised posterior portion in front of the intersegmental halfring behind the transverse ridge, brown; terminal ring of 2 occupying slightly more than its apical one- third, this spot narrowly prolonged forward in the median line; dorsal marks of 3—7 more deeply constricted subapically than in male, pointing basad, becoming successively broader from before backward, the broad basal portions of light ground colour most conspicuous and largest on 2—4, subinterrupted before their fu- sion with the dark apical rings; brown and yel- low areas of tergites 8—10 ill-defined, the yel- low predominating. Valves not surpassing seg- ment 10; cerci short, triangular and finely pointed, yellowish. Measurements: d abdomen + appendages 33.5— 38.2 mm, hind wing 22.8—25.0 mm, the holotype male being the largest specimen; © 32.5 mm and 24.0 mm, respectively. Lieftinckia kimminsi Lieftinck (fig. 18) Lieftinckia kimminsi Lieftinck, 1963: 537-541 (49 inclus. key), pl. 26 fig. 3 (6 wings), tfig. 17—18 (d struct.), 6 2 Bougainville I. No fresh material. This species is easily distinguished from its congeners by a combination of characters. It is chiefly remarkable for the well-defined bright blue head and thorax markings, and also by hav- ing a relatively large head (width across eyes and total body length in the ratio 10.2:100), which gives the insect a “facies” reminescent of certain species of Coeliccia. Shape of wings and neuration similar to Jairdi, but wings of the lat- ter are slightly narrower and less abruptly pe- tiolated (cf. Lieftinck, 1963: pl. 26, figs. 2 and 3). The pterostigma is rather like that of ramosa (fig. 13), distinctly less oblique, more regularly lozenge-shaped, than in lairdi. In both lairdi and kimminsi the posterior margin of the wing tips is strongly undulated, more markedly so than would appear from the published photo- graphs (loc. cit., same figs.). The male superior anal appendages are equal in length to, or a trifle longer than, segment 10. When viewed from within, the appendage shows a peculiar arma- ture, which is best shown in fig. 18. A broad, robust somewhat molar-like tubercle is placed about midway its whole length, at the base of the finger-like downpointing process; this tu- bercle is subrectangular in outline, with shal- | | lowly emarginate margin, its main portion being bluntly triangular and about equal in size to the | true apex of the appendage. In these respects | kimminsi differs from ramosa, in which the in- | terior process is considerably smaller, while the | true apex of the superior appendages is longer | and more slender (cf. figs. 21 and 22). Lieftinckia isabellae spec. nov. (figs. 11, 19) Material. — Solomon Is.: 3 d (two adults, one headless, one subadult), Santa Ysabel I., Maring Distr., Ta Matahi, 2.vii.1960, C. W. O’Brien. Holo- type d adult and one paratype & (ML), one paratype 3 (BISH). Stature more robust than lairdi and the larger-sized kimminsi, with a shorter and more compactly built synthorax, narrower wings, more normally lozenge-shaped pterostigma, lighter coloured end segments of abdomen, and quite differently shaped anal appendages. Male. — Labium yellowish, palpi ferrugi- nous, emargination of midlobe roundish, U- shaped, its depth less than ‘4 whole length of mentum. Labrum, mandibles (save the tips light ferruginous), anteclypeus and whole genal area, deep brownish black mixed with ferruginous, surface of all parts brightly shining. Postclypeus subhorizontal, its anterior 2/5 deep black, this mark widest at middle and convex behind, the anterior ridge sharply pronounced; colour otherwise bright chrome to orange (full col- oured male), whole surface shiny, the light col- our surpassing fronto-clypeal suture and con- tinued upward to form a complete broad trans- verse band of orange extending from eye to eye, occupying the frons as far dorsad as the rather swollen black antennal sockets. Rest of the head above, inclusive of the occipital ridge, unicol- oured dull velvet brownish black; rear of head otherwise ochreous. Antenna rather shorter than upper orbital line; scape short and cylin- drical, only little longer than its own diameter, black with slight greenish hue; pedicel much more slender, subequal in length to scape but about twice as long as its diameter, colour yel- lowish; flagellum indistinctly segmented, black. Main body of prothorax much shorter than in kimminsi and lairdi, hardly twice as long as its anterior lobe, the latter shaped similarly though much larger in size; pronotal tubercles weakly convex in side view, relatively larger and broad- LIEFTINCK: New Guinea and Solomon Islands Odonata 275 er than in the two species mentioned but of the same simple collar-like form, the side-edges narrowly rounded off. Synthorax distinctly shorter and more ex- panded than in both kimminsi and lairdi. Col- our-pattern a soft light rusty brown variegated with a mixture of delicate cream-yellow and pale green, the latter forming incomplete, ill-de- fined antehumeral bands tapering upward, sepa- rated above from a pair of almost conjoined jux- tahumeral patches of the same colour, one placed before, the other just behind the suture; metapleurae marked with two, almost complete light bands, both ill-defined but of the same tnt as the dorsal spots, separated from one another by an equally diffuse rusty brown band over the second suture. Whole ventral surface of thorax uniform pale ochreous, this tint turning to light green only along ventral metepimeral ridge. Legs throughout light rusty brown, as on most parts of the thorax, posterior femoral ridges finely black from near base to apex, the knees and extreme base of tibiae, distinctly ringed with dark brown, diffuse lighter brown annules placed slightly beyond halfway length of femora also discernible in two out of three males; colour of tibiae and tarsi as well as all bristles somewhat more obscured than on the femora; hind femur long, reaching back to very slightly over the apical border of segment 1, with 11—12 long bristles in outer row. Wings notably longer in proportion to the relatively short and very slender abdomen: fore wing when folded attaining base of segment 7; shape and neuration as in fig. 11; marginal un- dulations of apical portion quite distinct; veins almost black, membrane subhyaline. Arculus at Ax, in all six wings; M, arising at or a trifle dis- tal to the subnodus, Rs at Px,; anal bridge reaching the wing border at level of the prolon- gation of Ax, or sligthly distal to that point. Fore wing with 21—22, hind wing with 18—19 Px of first series. Area enclosed by C and R, posterior to pterostigma irregularly veined: very rarely with only a single cell-row, usually made up of more than 2 (1—6) duplicated cells (fig. 11). Abdomen slender, though relatively a little shorter than in both kimminsi and lairdi, from segment 7 on gradually somewhat expanded. Colour much as described for the other congen- ers: tergites bright yellow to dusky ochreous (depending on the state of maturity), but greater part of 2—6 much darker dorsally, brown to al- most black from end to end, the light parts forming more or less distinct basal rings, except 1 unmarked and basal one-third of 2 also re- maining yellow, its dorsal black mark pointing forward; 8—10 and anal appendages pale, lack- ing dark areas, only 7 occasionally obscured upon middle of tergite. Anal appendages short and compact, shaped as in fig. 19. Measurements: abdomen + appendages 33.8—35.2 mm, hind wing 22.5—24.5. Female unknown. Lieftinckia ramosa spec. nov. (figs. 13, 20—23) Material. — Solomon Is.: 1 d (holotype, adult, right hind wing and apex of left fore wing missing), San Jorge islet, off Santa Isabel, 26.1x.1965, “ Casuari- na forest, low herbage near stream”, Roy. Soc. Exped. Brit. Mus. 1966 (BM) [“small island to the SW of San- ta Isabel separated only by a small channel, but has a slightly different geological formation”, pers. comm. by Peter Ward, BM]. — 1 ¢ (adult, paratype), la- belled “Jdiocnemis inornata Selys Bougainville”, in R. Martin’s handwriting (ex MP, in ML). Male (holotype, San Jorge). — Labium dark brownish orange. Labrum, mandibles, genal area and anteclypeus obscurely reddish black, very shiny; postclypeus (with the anterior bor- der remaining black) bright shiny orange-ru- fous, as are also a pair of trapezoidal spots, one each side, connecting edges of postclypeus with inner orbital margin in front of antennal sock- ets, which are dull black; frons, antennae, ver- tex, epicranium and occipital region, all lustre- less deep black, only a small area next to the for- amen on postocciput somewhat lighter, reddish black. Antenna black, scape thick, squarish, the pedicel more slender, cylindrical, but only 1% times as long as scape; distalia missing. Prothorax, including anterior and posterior lobes, reddish black above, the sides throughout deep chrome; pronotal tubercles low, evenly convex; posterior lobe broad, not modified, de- pressed, almost straight behind, but distal bor- der distinctly upturned, with rounded side-an- les. ; Synthorax predominantly dark rufous inter- mingled with vinaceous brown, variegated with paler vinaceous and yellow areas which are no- where sharply outlined; black are: mesoprescu- tum, middorsal carina, margins of ante-alar tri- angles, and shiny black dots at all dorsal ridges inclusive of lateral sutures and posterior edge of 276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) 13 ae Pd Fig. 13. Wings of Lieftinckia ramosa spec. nov. (Bougainville). metepimeron. Mesepisterna with faint indica- tion of lighter antehumeral bands, which are broadening and somewhat better defined up- ward to form definitely yellow, subcircular dots; sides with some lighter colouring dorsally upon the mesepimeron and a wedge-shaped chrome-yellow streak, pointing upward, along mid portion of first suture, the latter not quite reaching metaspiracle; some yellow also on middle of mesinfraepisternum and posterior edge of metepimeron; ventral surface pale. Legs light rufous, but coxae paler and all fem- ora with ill-defined, broad, brownish rings and dark brown apices; all spines and bristles ru- fous. Shape and neuration of wings almost exactly as shown in fig. 13, for the male from Bougain- ville, but pterostigma a little more oblique and higher than in the type, its costal side distinctly shorter than the proximal, and distal side con- vex outwardly. Fore wings with 18, one hind wing with 17 Px of first series; arculus distinctly distal to Ax,. Pterostigma dark reddish brown. Abdomen very slender, terminal segments, from base of 8 to end of 10 gradually broad- ened, shaped much as in fig. 20. Colour mainly dark brown, segment 1 pale ferruginous above, the raised posterior border obscured and the sides also somewhat darkened; 2 obscurely brown with a diffuse yellowish basal annule and an indistinct stripe of the same colour along lower margin of tergite; 3—7 dark brown deep- ening gradually to almost black posteriorly, each showing low metallic lustre above and all with small but fairly distinct yellow basal rings, prolonged caudad alongside for a short distance, those on 3 and 7 narrowly interrupted by brown at the middorsal line; intersegmental rings obscured but on 3—6 preceded laterally by minute, ill-defined yellowish apical spots; dorsum of 8 lighter brown, its sides more broadly ferruginous; 9—10 and anal appendages entirely brownish orange to ferruginous, the swollen postero-lateral border of 10 with a black streak. Anal appendages as in fig. 20; apices of inner spur of superior pair as well as the incurved tips of the inferiors, black. Male (paratype, Bougainville). — Resembles the type in most respects, but differs slightly in body colour and structure of appendages, as fol- lows. Glossy surface of anterior parts of head uniform dark rufous instead of almost black, transverse clypeal band rather more orangish, less defined, and all the rest of head including antennae and undersurfaces, rufous, lacking dark areas. Pro- and synthorax as described for the type, except that there are no definite yellow areas on the mesepisterna, which instead bear a pair of complete, narrow and straight, cinna- mon-coloured antehumeral stripes not strongly contrasting against the dark orange-rufous ground; sides with faint indication of a greenish superior spot just below dorsal crest of mesepi- LIEFTINCK: New Guinea and Solomon Islands Odonata 277 Figs. 14—18. Platycnemididae. — 14, Male anal appendages of Salomocnemis gerdae spec. nov., dorsal and right lateral view; 15, Idem, ligula of male left lateral view and ventral view; 16, Idem, mesostigmal laminae of female; 17, Lieftinckia salomonis Kimmins, male anal appendages, dorsal and right lateral view; 18, L. kımminsı, left superior appendage, innerside. meron, also traces of light green on infraepister- nite and lower edges of metepimeron. Legs bad- ly damaged and broken. Wings (fig. 13), neuration brown. Fore wings with 18, hind wings 15—16 Px of first series; pterostigma brown. Abdomen shaped and coloured as in the type, except tergite 8 also showing traces of a latero- basal yellow spot. Anal appendages (fig. 23), very similar to those of the type, but lower branch of superiors slightly thicker and shorter, forming an angle of about 80° with the main branch (obtuse-angulate in the type), and inner tubercle about halfway length of superior ap- pendages a little less pronounced. Measurements: d abdomen + appendages 278 31.3 mm, hind wing 20.2 mm (San Jorge); 30.4 and 20.0 mm, respectively (Bougainville). Female unknown. This very distinct species has no near allies among its congeners inhabiting the Solomon chain but it approaches kimminsi most closely by having only a single row of cells between the veins C and R, beyond the pterostigma, and by having a branched apex of the superior append- ages. It differs, among other characters, from both kımminsi and lairdi by its more abruptly petiolated wings, less pointed wing tips, and the quite characteristically shaped and coloured head and thoracic markings. I am not quite certain about the correctness of the locality “Bougainville”, as written on the label by R. Martin himself. In fact several in- stances are known of Odonata in Martin’s col- lection whose reported habitats later proved to be erroneous. Lieftinckia spec. indet. Material. — Solomon Is.: 1 2 (subadult), San Cris- toval I., Wainoni, 22.vii.1965, Roy. Soc. Exped. Brit. Mus. 1966 (BM). This species is no doubt a near ally of L. salo- monis, but apparently quite distinct. Although the shape of the prothoracic hind lobe with its lateral blades and the peculiarly inflated meso- prescutum are developed similarly to those of salomonis, the mesoprescutal outgrowth in the present specimen is much narrower and sulcate medially, while the posterior lobes of the pro- thorax are subtriangular in outline. The colour- pattern also resembles that seen in salomonis; and, like that species, there are two cell-rows between the nervures C and R, beyond the pte- rostigma. This San Cristoval insect is, however, a good deal smaller in size than salomonis. By the absence of a topotypical male I prefer to leave this species unnamed. Lieftinckia spec. indet. Material. — Solomon Is.: 1 © (juvenile, left hind wing missing), San Cristoval I., Camp 2, 24.vii.1965, Roy. Soc. Exped. Brit. Mus. 1966 (BM). This © belongs to another undescribed spe- cies left without a name. It is immature and in rather poor condition. Posterior lobe of prothorax trapezoidal, rath- TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) er saddle-shaped lacking projecting side-lobes; no mesoprescutal outgrowths. Only a single row of cells between C and R, beyond ptero- stigma. Size small: abd. 31.5 mm approx., hind wing 22.5 mm. COENAGRIONIDAE Teinobasis Kirby, 1890 Teinobasis is a fairly large genus with over sixty named taxa and probably more than twen- ty still undescribed species and subspecies. The genus is wide-spread all over southern S. E. Asia, Micronesia, the Melanesian islands, and the northernmost parts of Australia. Only two or three outlying members, doubtfully gener- ically distinct from Teinobasis, are known from Malawi, a country on the verge of Ethiopia proper, and the Malgassian subregion (genus Seychellibasis Kennedy, 1920). At the time of summarizing the regional spe- cies of Teinobasis (Lieftinck, 1949), no members other than 7. aluensis Campion and rufithorax (Selys), had been known to occur in the Solo- mon Islands. On that last occasion, three more species were added to the list, but all were based on single specimens: two on immature females and the third on a male erroneously attributed to Papuagrion Ris, 1913. In the present paper all of them are revised, partly synonymized, and their structures more properly analysed. This has led to the recognition of 8 species (including one left without a name), of which the following 7 are believed to be validly named, viz., T. aluensis Campion, bradleyi Kimmins, chiono- pleura spec. nov., imitans spec. nov., obtusilin- gua spec. nov., rufithorax (Selys), and simulans spec. noy. As to the most widely distributed and best known rufithorax, this is only briefly men- tioned in the next pages and will not concern us here any further. All others are probably en- demic to the Solomons. Insufficient acquaintance with 7. aluensis must be held responsible for my erroneous as- signment of gurneyi to a new species of Papua- grion Ris, 1913. The latter is a Papuasian genus closely related to Teinobasis but with a more re- stricted range, comprising many, generally more sturdily built damselflies centred in New Guinea. Considering all characters used to sepa- rate these two genera (Lieftinck, 1932, 1935, 1937-38, 1949), and after re-examing the type- species, I arrived at the conclusion that 7. LiEFTINCK: New Guinea and Solomon Islands Odonata 279 19 Figs. 19—23. Male anal appendages of Lieftinckia species. — 19, L. isabellae, dorsal and right lateral view; 20, L. ramosa spec. nov., dorsal and right lateral view (San Jorge); 21, L. ramosa spec. nov., innerside of superior appendage (San Jorge); 22, Idem (Bougainville); 23, Idem, left lateral view (Bougainville). aluensis, emarginata and gurneyi should be re- united and placed in Teinobasıs, the last two be- ing obviously conspecific with alwensis, i.e. the earliest described member of this insular group. They were also compared with a few more re- cently discovered species, mentioned above and described below as new. They form a closely in- terrelated group differing markedly in details of structure and venation from Papuagrion (for references, see above). With the exception of the smaller-sized ones amongst the new taxa, two of them, simulans (New Georgia) and imitans (Guadalcanal), come very near aluensis and with much more material may even turn out to be only insular subspecies of the latter. As only few specimens are available for comparison, and no representatives of the aluensis cluster have so far been discovered in the intervening islands of the Solomons, questions of subspeciation and relationship cannot yet be answered. In the pre- sent case it seems best to keep all taxa apart as full species. It should be borne in mind that all of them exhibit a certain amount of variation in body colour and extent of markings, a well known disadvantage mainly depending on the insect’s stage of maturity, — in fact a real hand- icap frequently hampering easy recognition of similar-looking species of Teimobasis and allied genera! In the present context a few remarks are per- haps worth attention bearing upon the intricate and diversified structure of the end segments of the male abdomen, especially with regard to the conjunction and flexibility of their relation to one another. These sclerites, with their append- ages, are most complex in certain genera pres- 280 ently still united in the subfamily Pseudagrioni- nae (Davies & Tobin, 1984), a very large and heterogeneous group of genera. Some of the Old World members, like Pseudagrion and Ar- chibasis, contain generalized and easily recog- nizable forms, whereas others are more highly specialized, viz. Amphicnemis, Nesobasis, Pa- puagrion, Pericnemis, Teinobasis, and some an- nectent genera with fewer species. Regarding the last group, it may be remembered that I once redescribed and illustrated some species of Amphicnemis and Teinobasis occurring in the Philippine Islands. These had been character- ized already by Brauer (1868: 541—546), de Se- lys (1877: 114), and subsequently also by Need- ham & Gyger (1939). On two occasions (Lief- tinck, 1953: 250 and 1957: 161—170), I pointed out that the lower (inner) branches of the upper pair of appendages had been mistaken by all au- thors for the inferiors, possibly because the lat- ter are sometimes longer and stronger than the upper (main) branches of the superiors, OCCa- sionally abutting tightly on the major part of the inferior pair as well, even in the remarkable zy- gopteron Pericnemis stictica Selys. In that spe- cies the slender lower branch of the superiors is almost as strongly developed as the pincer-like upper portion. In all known genera of the Tei- nobasis alliance, the superior appendages are deeply divided and consist of two well-devel- oped branches. From dissections and pictures of these sclerites drawn in caudal view, it became quite clear that these processes are parts of a single appendage whose branches are firmly and immovably connected basally, forming together the true superior appendage. Consequently, the interpretation and terminology of the male gen- ital organs as given by Needham & Gyger (1939), are incorrrect. A further complication sometimes averting immediate recognition of the various compo- nents of the terminalia, is caused by the exis- tence of a median sclerite emerging from be- neath the overhanging ridge of the posterior border of the 10th tergite. This accessory scle- rite is usually placed in a vertical plane so as to protect the aperture and the more weakly scle- rotised membrane of the body wall. In the course of years, numerous drawings of that structure were published showing it in caudal (see e.g. Lieftinck, 1962, and the author’s papers (1932—1949), cited above. It is here termed “Breech-Block”; and though showing great di- versity in size and shape among species, its form seems to be quite stable specifically. Its function TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) is unknown, but in view of the complex form of | the female pronotal lobes and mesostigmal lami- nae, its flexibility is supposed to play an impor- tant role in the pre-copulatory process. Teinobasis aluensis Campion (figs. 24—26) Teinobasis aluensis Campion, 1924: 614, fig. 2 (6 app.), 6 Alu I. (Shortland Is. group). — Kimmins, 1934: 107 (type fixed, 2 topotype, not descr.). — Lieftinck, 1949: 343 (note on type). Teinobasis emarginata Lieftinck, 1949: 344—345, fig. 6(® proth.), © juv., Shortland Is. Syn. nov. Papuagrion gurneyi Lieftinck, 1949: 341—343, fig. 5 (6 app.). — 4 Bougainville I. Syn. nov. Material re-examined. — Solomon Is.: 1 d (adult), Alu, Solomon Is., C. M. Woodford 87—3, with D. E. Kimmins’s confirmative type label, holotype T. aluen- sis Campion (BM); 1 2 (juvenile), topotypical, det. D. E. Kimmins (BM). 1 © (juvenile), Shortland Is., Hisiai _ River, 22.x.1936, R. A. Lever, holotype T. emargina- ta Lieftinck (ML). — 1 d (adult), Bougainville I. 6.vili—16.x1.1944, A. B. Gurney, holotype Papua- grion gurneyi, Lieftinck (USNM). To the original description a few more details can be supplemented. Male (adult, holotype Alu I.). — Labrum broadly bordered with yellow-ochre rounda- bout its obscured centre. Face and frons entirely blue inclusive of the impressed area of the frons. Antennae pale (incomplete). Prothorax short, not modified. Synthorax with complete bronze- black middorsal band occupying the inner halves of the mesepisterna, beyond which the light non-metallic ground is discoloured and obscurated; the dorsum gradually acquires a lighter tint beyond the humeral suture, but the somewhat obscured bluish mesepimera are en- tirely without the distinct wedge-shaped bron- zy mark(s) seen in simulans and imitans; sides otherwise blue fading to yellowish underneath. Legs short, resembling Papuagrion in form; femoral bristles relatively strong, numbering 3, 4, —, respectively (posterior legs missing), slightly shorter than the interspaces; tarsal claws lacking a subapical inferior tooth. Wings much as described for gurneyı (loc.cit., 1949) and the other close allies; M, and Rs closely ap- proximated at origin, M, weakly curved, arısing very slightly before, Rs at the subnodus, both veins remaining separated thereafter. Ab enter- ing the posterior margin well beyond Ac; me- dio-anal link entire; three postquadrangular an- tenodal cells. All wings with 15 Px of first se- ries. Pterostigma as in simulans. | | LIEFTINCK: New Guinea and Solomon Islands Odonata 281 sup. app. (lower Ze branch) Figs. 24—33. Teinobasis — 24, T. aluensis Campion, male anal appendages, left lateral and dorsal view (type of T. gurneyi); 25, Idem, caudal view; 26, T. aluensis Campion, hind margin of prothorax of female (type of 7. emarginata); 27, T. simulans spec. nov., male anal appendages, dorsal view (New Georgia); 28, Idem, left lateral view; 29, Idem, caudal view; 30, T. imitans spec. nov., male anal appendages, dorsal view (Guadalcanal I.); 31, Idem, left lateral view; 32, Idem, caudal view; 33, 7. imitans spec. nov., prothorax of female, dorsal view (Gua- dalcanal I.). 282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) Abdominal tergite 10 in caudal view shaped much as shown for the type of gurneyi (fig. 25) and imitans spec. nov. (fig. 32), the vertical sur- face of the raised upper part of the body-wall slightly convex, black in colour, but its mid-api- cal process bright yellow (deep black in simu- lans), at first directed obliquely caudad, then strongly hooked, recurved, with bilobed apex. Anal appendages equal in length to segment 10; colour yellow; superior pair relatively short and broad in dorsal view, slightly but distinctly S-shaped in side view, as shown for gurney) (fig. 24), Measurements: abdomen + appendages 48 mm, hind wing 31.5 mm. Female (juvenile, “red” colour phase, Alu I.). — This (undescribed) teneral specimen was ex- amined by me at the British Museum (Natural History) after publication of T. emarginata, with which it agrees in most respects, especially in the venation. Prothorax similar to emargina- ta, except that the deeply divided lamellae of the posterior lobe are somewhat more drawn out at the inner edges, so as to resemble the adult fe- male of imitans (fig. 33) more closely; in emar- ginata the lobes are more depressed and shrunken owing to the teneral condition of the sclerite. Metallic green marks covering the tho- racic dorsum in the form of ill-defined bands occupying the inner halves of the mesepister- nites. Its measurements are: abdomen 41.5 mm, hind wing 30 mm. Teinobasis simulans spec. nov. (figs. 27—29) Material. — Solomon Is.: 1 6 (adult), New Georgia I., Bareki River, 30.viii.1965, Roy. Soc. Exped. Brit. Mus. 1966—1. Holotype (BM). Male (adult). — Labium yellow; basal half of labrum (including the central pit) glossy brown- ish black, its distal half sharply defined bright ochreous; mandible-bases and genae dark olive, anteclypeus more brownish olive, postclypeus dark brown. Frons with well pronounced, al- most rectangulate transverse anterior crest, the smooth vertical surface dark olive; upper sur- face of frons, antennal sockets and all remaining dorsal parts of head dull greenish bronzy black with some traces of brown just in front of the ocelli and along occipital crest; occipital area and rear of the head dirty orangish. Scape and pedicel of antenna yellowish, the latter about 1/2 times length of scape, its apex brown; dista- | lia black. Pronotum dark brown, lower portion of tu- bercles fading to rusty brown, the latter low and but little convex; sides including those of poste- rior lobe rusty brown, the lobe short and broad, longest at middle, somewhat upturned, its sur- face convex anteriorly; posterior border in dor- sal view throughout evenly and but slightly convex, the side lobes small, not protuberant and rounded. Dorsum of synthorax as far down as a little before humeral suture and including ante-alar triangles, black with low but distinct metallic green lustre, a hairline at the humeral suture re- maining rusty brown; mesepimeron rusty brown, its lower (anterior) half marked with a wedge-shaped brown spot pointing upward and ill-limited above, posterior portion yellowish green, as are also the remaining parts of the sides and ventral surface of thorax; mesinfra- episternites rusty brown with a subcircular blackish central dot low down. Coxae and trochanters of legs light green; legs otherwise pale ochreous; but outer faces of all femora with two brown stripes partly fused together beyond halfway length; extreme bases and carinae of tibiae as well as all bristles, brown. Hind femora with 4—5 strong bristles in outer row, all a little shorter than the inter- spaces; tarsal claws lacking an inferior subapical tooth, or a rudiment of a tooth only at the fore tarsi. Wings narrow, venation brown. Ac situated much nearer Ax, than Ax,, its length a little less than half its distance from proximal side of qua- drilateral, which is scarcely longer than Ac it- self; Ab enters the margin well distal to Ac at a distance of about twice the length of Ac. Arcu- lus at Ax, in all wings. Quadrilateral in fore wing with proximal and costal sides of equal length, in hind wing costal side markedly longer than proximal side. Medio-anal link entire; three subquadrangular antenodal cells. M, mod- erately curved at origin, arising sligthly before the subnodus (least so in fore wings), Rs at the subnodus, both veins very closely approximated (just not fused together), from their meeting point to almost as far as Px,, thence gradually diverging. Broken course of M, and Cu, dissi- milar: fractured portion of M beginning beyond halfway its length, that of Cu, much before that level, i.e. at level of Px,-Px,. Fore wing with 15—16, hind wing with 14—15 Px of first se- ries. Pterostigma of small size, braced, lozenge- LIEFTINCK: New Guinea and Solomon Islands Odonata 283 shaped but very oblique, a little longer than high, its proximal side only a trifle more oblique than the distal one; colour cinnamon, darker brown centrally. Abdomen long and slender, basal and termi- nal segments moderately expanded. Segment 1 light blue-green, distal one-fourth of tergite with crescent-shaped transverse brown mark and traces of rusty brown in front of the latter; dorsum of 2 metallic green from base to apex, this mark very slightly expanded before the end, the sides blue-green; succeeding segments for the greater part brownish black with slight bronzy reflections, 3—7 each with pair of small, greenish yellow basal spots forming dorso- lateral annules narrowly interrupted from be- hind in the median line; sides of 8 with a yel- lowish stripe, broadest at either end bordering lower margin of tergite; 9 with transverse basal stripe of yellow alongside; 10 with similar stripes basally, the sides as well as the apical 1/3 of the upper surface of this tergite likewise brownish yellow. Intersegmental rings of 8—9 and 9—10 pale. Configuration of 10th tergite and anal appendages as shown in figs. 27—29; colour brownish yellow, including the midapi- cal process of tergite 10; lower branch of supe- rior appendages ending in a transverse, smooth, shining black ridge bearing minute erect light hairs; fringes on the inside at tips of superiors and long tufts at apices of inferior appendages, all pale golden yellow. Measurements: d abdomen + appendages 46.6 mm, hind wing 29.0 mm, pterostigma 0.5 mm. | Female insular topotype, unknown. Teinobasis obtusilingua spec. nov. (figs. 37—39) Material. — Solomon Is.: 2 ¢ (adult, one in frag- ments), San Cristoval I., Huni River, mouth Camp 4, 14.vin.1965, “in and around small pools in Casuarina forest on basic soils”. Roy. Soc. Exped. Brit. Mus. 1966-1. The perfect specimen is the holotype (BM), the fragmentary one a paratype (ML). Male (adult, holotype). — Labium yellowish; labrum almost wholly black, only anterior nar- rowly lined with yellow. Mandible-bases yel- lowish with a large central spot of brown; genae and anteclypeus dirty brownish, postclypeus glossy brownish black. Frons shaped as de- scribed for chionopleura, colour deep black with faint bronze reflections, the sulcus between cly- peus and frons brown. Head otherwise dull bronze-black, save for a pair of small, oblique pale streaks, one either side, between base of an- tenna and lateral ocellus; occipital ridge also somewhat lighter. Rear of head apparently brownish black or black except anteriorly a tri- angular yellow mark at level of submentum. Antennal scape very short, black, at most one- third as long as pedicel, which is yellowish streaked with brown, flagellars long and slen- der, black. Prothorax including posterior lobe, bronze- black, the main portion bearing a broad x- shaped mark occupying the whole dorsum, sides greenish yellow; posterior lobe simple, de- pressed, much broader than long, surface con- vex, hind border evenly and broadly rounded, the side-angles narrow, not projecting, round- ed. Dorsum of synthorax, as far down as the hu- meral suture and including ante-alar triangles, black with distinct bronze and coppery reflec- tions; mesepimeron pale rust-coloured, marked with an irregular bronze-black band that covers the lower 2/3 of the area but dorsally narrows abruptly to a stripe extending upward along the impressed part of the first suture, the latter curving round the black dorsal crest so as to al- most meet an elongate blackish streak along up- per end of second suture; remaining parts of side and ventral surface unicoloured greenish yellow; mesinfraepisternites more rust-col- oured, with a large subsquarish patch of bronze-black covering most of that sclerite. Legs pale ochreous, the coxae mixed with green; outer faces of all femora and the knees narrowly, with complete thick dark brown stripe; four strong dark brown bristles in outer row and five in inner row of hind femora, all shorter than the interspaces. Tibiae and tarsi un- marked; tarsal claws simple, or with the merest trace of an inferior subapical tooth. Wings narrow, shaped as in bradleyi, the apices slightly more pointed. Ac much nearer Ax, than Ax,, only about one-third as long as its distance from proximal side of quadrilateral, which is nearly twice as long as Ac itself; Ab enters the margin well distal to Ac at a point away from it for about twice the length of Ac. Arculus distinctly distal to Ax, in all wings. Quadrilateral longer than in bradleyi, its costal side in fore wing longer than proximal side, in hind wing more markedly so. Medio-anal link entire; three postquadrangular antenodal cells. Origin of M, and Rs close together, M; moder- ately curved at origin, arising a trifle before sub- nodus in fore wings, at the latter in hinder pair; 284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) NN NN Figs. 34—43. Teinobasis. — 34, T. bradleyi Kimmins, male anal appendages, dorsal view (Bougainville I.); 35. Idem, left lateral view; 36, Idem, caudal view; 37, T. obtusilingua spec. nov., male anal appendages, dorsal view (San Cristoval I.); 38, Idem, left lateral view; 39, Idem, caudal view; 40, T. bradleyi Kimmins, thorax; 41, 7. chionopleura spec. nov., anal appendages, dorsal view (holotype male, Sandfly I.); 42, Idem, left lateral view; 43, Idem, caudal view. LiEFTINcK: New Guinea and Solomon Islands Odonata Rs arıses a little beyond the subnodus, while both veins gradually diverge immediately after their meeting point (in right hind wing they are fused together for a short distance!). Broken course of M, and Cu, as described for bradleyi. Fore wing with 13—14, hind wing with 12 Px of first series. Pterostigma of small size, long and narrow, braced, much smaller than under- lying cell and almost twice as long as high, sides oblique but proximal side distinctly more oblique than distal side; colour deep black. Abdomen long and slender, shaped much as in bradleyi; tergite 1 yellow-green with a trape- zoidal bronze-black marking occupying most of the dorsum and broadest posteriorly; 2 with the dorsum bronze-black from base to apex, this mark a little constricted at extreme base and hardly expanded subapically, the sides possibly greenish in life; succeeding segments for the greater part bronze-black, 3—6 or 3—7 each with narrow yellowish basal annules which suc- cessively become more broadly interrupted and placed more laterad posteriorly; 8—10 appar- ently unmarked, but intersegmental rings of ter- minal segments yellowish. Configuration of 10th tergite and anal ap- pendages as in figs. 37—39; colour black, except the dorso-apical ridge, the thickened posterior border of last tergite, and most of the inferior pair of appendages, which are light brown; sub- apical hair-fringe springing from inner faces of superior appendages, also pale. Measurements: d (holotype) 37.0 mm, hind wing 21.8 mm, pterostigma 0.5 mm. Female unknown. Teinobasis chionopleura spec. nov. (figs. 4143) Material. — Solomon Is.: 1 & (adult, holotype), Florida Is., Ngela group, Sandfly Island, Takavali, 22.x11.1965, “common in forest”, R. Slooff (ML); 1 3 (adult, discoloured), Florida Is., Small Ngela I., Dende, 18.1x.1960, C. W. O’Brien (ex BISH, ML); 1 3 (adult, head and part of abdomen missing), Florida Is., Big Florida I., 28.x1.1944, H. E. Milliron (BISH). Male (adult, Sandfly I.). — Whole labium, in- cluding submentum and all parts in front of the foramen, light chrome as are also the genae as far back as the base of squamae, only the mandi- bles tipped with black. Labrum, clypeus and a transverse band in front of frons connecting the eyes, which reaches up as far as the antennal bases and including the antennal sockets, all 285 unicoloured orange-chrome. Frontal ridge weakly developed, obtuse-angulate, surface of frons uneven but lacking transverse wrinkles. All the rest of head behind the frontal ridge, en- tirely dull greenish black to black without any light markings, the rear also wholly black but postoccipital lobes with a pair of bluish pruines- cent patches. Scape and pedicel of antenna of the usual shape, short and subequal in length, both obscured but apex of scape yellow-ringed. Pro- and synthorax entirely metallic bronzy black, all sclerites (except upper portions of meso- and metepimera) thickly overlaid with palest blue pruinescence, leaving only traces of the orangish ground colour at dorsal ends of hu- meral and first lateral sutures, and posterior edge of metepimeron. Coxae, trochanters and extreme bases of fem- ora, black, these parts pruinescent blue all over; legs otherwise light orange-yellow, the tips of femora narrowly obscured and all bristles black; femoral bristles slightly longer than the inter- spaces, numbering 2 + 2, 6—7, and 6, on the three pairs of legs; tibial bristles 5, followed by a row of 6—7 much shorter ones at fore tibiae, 4—5 at mid tibiae, and 5—7 at hinder pair of same; tarsal claws without any indication of a subapical tooth. Wings hyaline, neuration as for genus, but quadrilaterals rather long: proximal and costal sides subequally long in fore wings, but proxi- mal side only about half as long as costal side in hind wings, the costal and distal sides in the lat- ter being subequal in length. Arculus at Ax,; Ac much nearer Ax, than Ax,; Ab entering the wing margin a little less than 112 times the length of Ac itself beyond it. Three postqua- drangular antenodal cells; M, arises well before, Rs at the subnodus, base of M; normally curved and both veins well separated immediately after their meeting point. M, and Cu, of normal great length, marginal convexities at end of veins hardly indicated. Fore wing with 16, hinder pair with 15 Px of first series. Pterostigma distinctly oblique, lozenge-shaped but parallel-sided, 1% times longer than high; colour deep black finely bordered with yellow. Only a single row of 4— 5 cells between C and R, posterior to pterostig- ma. Abdomen long and slender, from end of seg- ment 6 to base of 10 slightly expanded, the latter parallel-side. Ground colour yellowish, but all tergites with predominantly light brown mark- ings, this colour gradually deepening to dark brown and bronze-black toward end of abdo- 286 men, 8—10 (except laterally) mostly black. Ter- gite 1 with diffuse brown dorsal mark broaden- ing toward apex forming an ill-defined dark ter- minal ring, the sides bright greenish yellow; a complete dark mark also on dorsum of 2, deep- est in colour and broadest on basal half but leav- ing tiny yellow streaks on either side at base, the sides also greenish yellow; dorso-lateral bands on 3—7 suddenly abbreviated in front, so as to save a pair of small yellow spots at the bases of each, the latter being largest on 5—7 and co- alescent with distinct citron-yellow lines bor- dering ventral margin of tergites; 2—6 more- over with fine deep black intersegmental an- nules restricted to the dorsum, those of 8—10 bright yellow; sides, distal one-third of tergite 10 as well as the vertical mid-caudal process of the latter, ochreous. Anal appendages (inclusive of the 10th apical plate) ochreous, lower branches of superior ap- pendages and the incurved apices of main branch, black-tipped; inner faces of superiors clothed densely with softish pale pubescence. Measurements: abdomen + appendages 43 mm; hind wing 27.3 mm. Male (adult, Small Nggela I.). — This para- type is the 2nd complete d whose characters correspond almost exactly with those of the type, except that there are minute traces only of pruinescence, the thoracic sclerites being wholly deep black with low bronze reflections, save the posterior edges of metepimeron and the post- sternal surface, which are light ochreous. The legs also are coloured as in the type, only the tarsi being obscured. Size slightly larger: abdomen + appendages 46.0 mm, hind wing 28.4 mm. Male (adult, incomplete, Big Florida I.). — Quite similar to the last-mentioned male. How- ever, there is an interesting point of difference in its wing venation: the nervure Ac is placed further distad in both hind wings, i.e. at a level only little away from Ax, causing the veins Ac and Ab to meet the posterior border of the wing at one point, which is unusual for Teinobasis. In all specimens the medio-anal link is entire. Female unknown. This new species is distinctly more slenderly built and has a somewhat larger head, than the members of the 7. aluensis group, though all are of about the same large size. Further peculiari- TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) ties are found in the structure of the terminal genital organs by which chionopleura differs from its congeners in the Solomons. In caudal view, the raised vertical portion of the 10th ter- gite (fig. 43), does not seem to bear a prolonged median process but is reduced to a short trans- verse plate hardly visible in dorsal aspect. The black-tipped upcurved and pointed lower branches of the superior appendages are placed inward to a pair of shorter, blunt and pale-col- oured, upper branches of the inferior append- ages, both of these being well visible in side view (fig. 42), but hidden from view when looked at from above (fig. 41). Teinobasis imitans spec. nov. (figs. 30—33) Material. — Solomon Is.: 1 d (adult holotype), Guadalcanal I., Honiara, 150 m, 2.v.1964, J. Sedlacek (ex BISH, ML). 1 © (adult), Guadalcanal I., Honiara, . 17.1x.1965, “at house lights”, Roy. Soc. Exped. Brit. Mus. 1966 (BM). Closely allied to both 7. aluensis and simulans spec. nov. Male (holotype, adult, abdominal segments 4—7 missing). — Labium, mandible-bases, and genae as far up along orbital lines as base of an- tennae, obscurely greenish yellow (possibly green or blue in life), mandibles tipped with deep black; labrum bright chrome gradually changing into brown at extreme base, the boundary almost straight but ill-limited. Ante- and postclypeus black with bronze lustre, sur- face of anteclypeus uneven, of postclypeus fine- ly transversely striated. All the rest of head, in- clusive of postocular lobes and occipital ridge, almost lustreless dark bronzy black; transverse frontal crest distinct, subacute; rear of the head, except some diffuse obscuration on inner sur- face, bright greenish yellow. Antennal sockets somewhat swollen, scape short and thick, pedi- cel much slenderer, about 1/2 times length of scape, both obscurely yellow. Prothorax dirty ochreous mottled with dark brown on pronotum and centre of pleurae, the anterior lobe suberect, strongly black-rimmed in front; posterior lobe broad, collar shaped, de- pressed, its hind margin somewhat undulated, the side-edges rounded, whole surface dark me- tallic green. Synthorax robust, shaped as in aluensis and simulans spec. nov.; centre of mesostigma lamina and whole mesonotum with the ante-alar triangles, dark metallic green to as LIEFTINCK: New Guinea and Solomon Islands Odonata 287 far as the humeral suture, which is indicated by a yellow hair-line; mesepimeron and mesinfra- episternum beyond that level, mainly light blue, the former bearing two distinct patches of dark metallic green of unequal size, separated from each other by the pale ground colour: a diffuse, oblique zone of a more ferruginous tint. Meta- pleurae similarly coloured but changing to (or mixed with) clear blue upon middle of metepis- ternum and to yellow further down, the whole ventral surface of thorax also unmarked. Legs almost entirely yellow, only outer faces of coxae light blue and those ot femora slightly more orangish with the knees a shade darker, the yellow at the tibiae conspicuous; femoral bristles robust, brown, 2—4—5, respectively, little shorter than the interspaces; tibial bristles shorter than interspaces, fore tibiae with four longer bristles followed by a row of ca. 10 thin- ner ones (setae) in outer row, 4—5 strong ones on mid and hind tibiae. Tarsal claws indistinctly toothed: rudimentary or obsolete on all legs. Wings long, distinctly petiolated as fas as Ac, much as in aluensis, i.e. more markedly broad- ened than in the smaller-sized regional Teinoba- sis, but again broader than in the close ally simu- lans; venation brown. Arculus at Ax, in all wings; position of Ac and Ab as in the other species: Ab entering margin at a point in line with proximal side of quadrilateral, the latter in fore wing only little longer than, in hind wing almost twice as long as, costal side. Medio-anal link entire; three postquadrangular antenodal cells. M; arises well before, Rs at the subnodus, the former moderately curved, both veins well separated after their subnodal meeting point. M, and Cu, of great length, the broken course of M, beginning at Px, ; in fore wings, at Px, 9 in hinder pair, and of Cu, at Px, ;. Fore wing with 16, hind wing with 15 Px of first series. Pterostigma small, oblique, much longer than high, lozenge-shaped but subparallel-sided, braced and much smaller than underlying cell; colour brown narrowly lined with yellow along proximal and costal sides. Between C and R, posterior to pterostigma 1—2 divided cells about halfway length of space (undivided in right hind wing only). Abdomen (incomplete), ground colour of ter- gites 1—2 blue, on succeeding segments dirty ochreous; 1 with complete subtriangular dorsal mark of metallic green pointing basad, sides broadly blue; dorsum of 2 metallic green from base to apex, leaving only a pair of narrow sub- interrupted blue annules at extreme base, the lower half of sides yellow; dorsum of 3—7 ap- parently similarly banded, but losing gradually their metallic gloss and becoming broader and darker toward end of abdomen, only at base of 3 a pair of semicircular yellow spots confluent with the pale ground colour at the sides; 8—10 almost wholly black, but intersegmental mem- branes of 8—9, 9—10 and 10—sup.app., stand- ing out clearly as bright ochreous dorsal an- nules, the swollen dorsolateral ridge at apex of tergite 10 also yellow. Anal appendages (figs. 30—32) subequal in length to segment 10, shaped very similar in principle to those of aluensis, its synonym gur- neyi, and simulans. Dorsal (main) branch of su- perior appendages black above and black- tipped, yellow and rather hollowed out within, each provided with tuft of long pale golden hairs (resembling szmulans); inferior appendages also yellow above and densely light-haired, obscured ventrally. Female (adult, topotype). — Resembles the male very closely in most respects, differing on- ly in colour design of body and legs, as follows. Labium, upper mouth-parts, face and frons as well as first two antennal segments, all light ochreous, but at mandibles and genae slightly mixed with olive, the labrum being more bright- ly orange. Rest of head above dull bronze- black, sharply contrasting with the greenish yel- low rear. Antennal pedicel dark-ringed apically, the distalia black. Prothorax (fig. 33), reddish ochreous, mar- gins of posterior lobe obscured, the latter hardly raised; laminae mesostigmales transverse, carry- ing light brown, erect, ear-like lobes. Synthorax much as in male, bronze-green dorsally to just before humeral suture, the suture itself pale, the bronzy tint including ante-alar triangles, mid- ventral and axillary sclerites; colour at sides and underneath, all dirty ochreous. Brownish patch upon shoulder area (mesepimeron) very diffuse, with slight metallic green gloss. Legs totally yellow save outer faces of coxae blue and those of femora orangish, armature as in male. Claws with rudiment only of an inner subapical tooth. Wings well petiolated, with brown neuration; medio-anal link entire though slightly fractured in left hind wing. Wing tips relatively obtuse; all wings with 15 Px of first series. Pterostigma as in male, light brown sur- rounded by yellow; area between C an R, distal to pterostigma exactly as in male. Abdomen yellow-brown gradually darkening 288 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) toward end; tergites 2—7 indistinctly obscured apically, 3—8 moreover with very narrow ill- defined yellowish basal rings interrupted by the yellow-brown dorsal colouring; intersegmental membranes dark, except those between 7—8 and 8—9, yellow; tergites 8—10 metallic green- ish black, the lower half of sides pale. Genital valves not surpassing apex of segment 10, carry- ing a row of minute, blunt marginal serrula- tions; cerci obscured, shorter than segment 10, divergent, pyramidal, tips acute. Measurements: d abdomen + appendages 47 mm (approx.!), hind wing 31.5 mm; © abd. 46.0 mm, hind wing 32.6 mm, pterostigma 0.6 mm. This is probably the closest relative of aluen- sis. Apart from minor colour features of less im- portance, male imitans differs from aluensis in the following points: (1), the whole posterior surface of the vertical process (or “breech- block”), at the apex of the 10th tergite, is deep black (bright yellow in aluensis); (2), the outer faces of the main (upper) branch of the sup. app. and their lower branches as well, are also black, the latter being very shiny; (3), the appendages are longer and more slender, less hollowed out within, than in aluensis; (4) the inf. apps. of im- itans are obscured ventrally but yellow on the inside and clothed densely with pale pubes- cence, the tip with the long fringe of pale golden hairs directed inward (only partly shown in fig. 31). In both species the median process of the 10th tergite is broad and somewhat flattened when viewed from above, but strongly hooked and downcurved in profile view (cf. figs. 30 and ID): Teinobasis bradleyi Kimmins (figs. 3436, 40) Teinobasis bradley: Kimmins, 1957: 315—316, tfigs. 2 A—C (3 thorax & anal apps.). — d Guadalcanal I.; Kimmins, 1970: 183 (8 lectotype selected). Material. — Solomon Is.: 2 d (adult, one imper- fect), Guadalcanal I., Tapenanje, 10—23.xii.1953, lectotype and paralectotype (BM). — 2 6 (adult), Ma- laita I., Ngwaiau, 1500 ft., 10.x.1967, “in forest”, R. Sloof (ML). — 1 d (adult), Santa Ysabel I., Maring dist., Te natahi, 2.vii.1960, C. W. O’Brien (ex BISH, ML). — 1 g (adult), Bougainville I., Buin, 2.1.1970, R. Straatman (ML); 2 6, 1 ® (adult), Bougainville I., Buin town, 15—16.v.1975, “along small stream”, H. R. Wimmer (USNM), and Bougainville I., 16.xi.1944, “ex jungle by stream” (one pair), A. B. Gurney (USNM); 1 ® (juvenile, red), Bougainville I., Buin, 4.v1.1956, J. L. Gressit (BISH). Male (adult, Tapenanje). — The following | colour and structural notes are additional to those given for the lectotype, subsequently se- lected by Kimmins for one of his specimens from Guadalcanal. Differs from T. obtusilingua spec. nov.: ante- rior margin of labrum more broadly bordered with chrome. Face and frons in front, bright blue. Thoracic pattern wholly different, as shown by Kimmins (loc. cit., tfig. 2 A) and in our fig. 40. Inner faces of sup. anal app. beset along whole length with long pale hairs (in ob- tusilingua only at apices); appendages very simi- lar in the two, but mid-apical process of tergite 10 broader and tapering to a blunt point, deep black, as shown in fig. 34 (see also Kimmin’s tfig. 2 C). From T. aluensis, the present species differs in that the vein M, arises before, and Rs at the subnodus, but this character may be unstable; aluensis is also a somewhat more ro- bustly built and larger species than bradley:, the type of the latter measuring abdomen + ap- pendages 39 mm, hind wing 26 mm. The “slight undulations of the wing apices”, mentioned by Kimmins, apply to the prominent posterior bor- der at the end of the main veins, not to the api- cal undulations, as seen in a number of regional platycnemidids. Female. — No insular topotypes are avail- able. Close comparisons made between typical bradleyi from Guadalcanal and all specimens collected in the island chain of Malaita, Santa Ysabel, and Bougainville, have led to the con- clusion that all these populations belong to a single species which, accordingly, seems to have a much wider distribution than was thought at the outset of this investigation. All agree in their wing venation, the scheme of coloration, and their genital structure, only the body size and details of colour having been noticed. The next descriptions and drawings are based on the best preserved and fully mature specimens from Bougainville, which also include a well-col- oured female from that most westerly situated island of the chain. Male (adult, Bougainville). — Labium palest yellow, labrum glossy black, the anterior bor- der, except laterally, yellow, a vestige of a yel- low spot also on either side at extreme base; mandible-bases light blue, the distal one-third brownish black; anteclypeus light blue, postely- LIEFTINCK: New Guinea and Solomon Islands Odonata 289 peus glossy black; entire genal area and ante- frons as far as the transverse ridge and insertion point of antennae light blue, this colour extend- ing posteriorly all along orbital margin to be- yond base of maxillae; remainder of frons and all the rest of head including the occiput, black. Frons and ocellar area with low bronze reflex, behind which the interocular and occipital re- gion with more distinct dark metallic green lustre. A tiny oblique yellow spot just visible between antenna and lateral ocellus. Occipital lobes slightly pruinescent blue, an oval patch of blue pruinescence placed more rearward also upon centre of occiput. Cervix yellow. Whole dorsum of prothorax black with slight bronze sheen, the sides light blue; pronotal tu- bercles low, evenly convex; posterior lobe sim- ple, rather crescent-shaped, surface convex and directed straight back, about twice as long as the side lobes, which are rounded off; whole border evenly rounded, hardly swollen or upturned. Dorsum and upper part of sides of synthorax, black, as shown in fig. 40, for the rest all vivid light blue, as described for the type by Kimmins (1957, fig. 2 A). All dorsal ridges and humeral suture with the exception of a vestigial spot of blue at either end of the latter (one to the inside, the second at the dorsal ridge of suture), like- wise black; inner portions of black mesepisterna with distinct metallic green lustre, the remain- der deep black; a sharply contrasting brown hair line laterally, and a pair of posterior dark spots on the poststernum. Coxae and trochanters of legs blue; femora pale greenish blue with complete, thick, dark brown exterior stripe broadening to an apical ring at the knees; tibiae yellow-brown tipped with ochreous; tarsi bright orange-chrome. All spines and bristles black; femoral bristles as long as the interspaces, 3, 4, 4, respectively; tib- ial bristles longer, also equalling interspaces, 5, 4, 5, respectively; tarsal claws lacking an inferi- or tooth, the tips obscured. Wings narrow, neuration much as in all other regional congeners; membrane hyaline, veins black. Ac situated much nearer Ax, than Ax,, only half as long as its distance from the qua- drilateral, which is slightly less than twice as long as Ac itself; Ab enters the margin well dis- tal to Ac, for fully twice the length of Ac. Arcu- lus at Ax, in fore wings, very little beyond that level in hinder pair. Quadrilateral of fore wing with proximal side little shorter than costal side, in hind wing costal side markedly longer, twice as long as proximal side and subequal to distal side. Medio-anal link entire; three postqua- drangular antenodal cells. M; and Rs closely ap- proximated at origin, base of M, but little curved, arising slightly before, Rs slightly after the subnodus, both veins well separated distal to their meeting point. Broken course of M, and Cu, dissimilar: fractured portion of M, begin- ning far distad, i.e. at level of Px,, in both pairs of wings, of Cu, at level of Px,, both portions in the latter subequally long. Fore wing with 13— 14, hind wing with 13 Px of first series. Pteros- tigma small, almost twice as long as high, oblique but almost parallel-sided, surmounting exactly one much larger cell; colour brownish black, costal and anal sides bordered with ex- tremely fine pale hair-line. Abdomen very thin and slender, basal seg- ments more expanded than terminal ones; sides of tergites 1—2 largely blue, dorsum of both with complete metallic greenish black band broadening markedly toward end though, after a subapical constriction, forming almost com- plete dark terminal annules, the intersegmental membrane 1—2 blue; dorsum and most of the tergal sides 3—7 occupied by well defined bronze-black marks from base to apex, leaving only vestigial paired yellowish spots at extreme base, the dark bands broadening somewhat pos- teriorly, the rest of the tergites bright ochreous in colour. Tergite 8 with the dorso-lateral mark hardly broadening posteriorly, on 9 almost straight-lined, and the one on 10 leaving only a triangular spot along lower border. All interseg- mental membranes dorsally as well as the ster- nites, bright chrome yellow. Configuration of 10th tergite and anal appendages as in figs. 34— 36, colour black or almost so, only inner shelves of main branch of inferior appendages yellow- ish, the hairs fringing inner faces and apices of upper branch of superior appendages distinctly white. Female (adult, Bougainville). — This was col- lected simultaneously with the male and corre- sponds closely with the latter in stature and ve- nation. Here follows a comparative description. Labium bright yellow. Labrum black only at middle along base, all the rest of it vivid orange. Anteclypeus obscured, dullish; postclypeus contrastingly deep glossy black, except laterally and on the mandible-bases which are coloured a dirty greenish ochreous, yet acquiring a more vividly green tint on the genal area and forming a broad colour band also on the antefrons as far up as the antennal sockets and frontal ridge; 290 first antennal segments partly obscured. Whole less shiny than the black postclypeus, light marks restricted to a pair of tiny oblique specks placed about midway between lateral ocelli and antennae; a thick transverse bar upon the occip- ital plate at some distance behind lateral ocelli, also pale. Rear of the head deep black, the pale genal area extending upward along eye-border as far as base of labial hinge, the black area thin- ly pruinescent. Prothorax ferruginous brown, its sides over- laid with bluish pruinescence; posterior lobe larger and more raised than in male, median portion dome-shaped in caudal view, its border broadened a little on either side of middle to form slightly forwardly curved, rounded lobes, which are convex anteriorly, hollowed out pos- teriorly. Synthorax marked exactly as in male, except that the dorsolateral parts are ferrugi- nous instead of black, only the middorsal carina and a complete parallel-sided band attached to it on either side, metallic green, this median band almost equal in width to each of the mesepister- nal (antehumeral) halves; remainder of thoracic sides as well as the ventral surface throughout pale blue, slightly pruinescent lower down. Legs coloured as in male, i.e. vivid light chrome, the bases of all coxae blue, a complete though narrow black stripe at outer faces of all femora, these stripes broadening at end to form distinct apical annules; bristles black. Shape of wings and neuration as in male, the relatively long pte- rostigma light brown finely surrounded with yellow. Px of first series 15 in fore wings, 13 in hinder pair. Abdomen comparatively slender, the terminal segments but little expanded. Ground colour greenish yellow, all segments marked with bronze-green dorsal bands, complete on tergites 1—2, both broadening toward end, those on 3—7 more abruptly narrowed basally and ex- panded apically, the small yellow basal annules finely interrupted in the median line; colour of apical tergites obscured, broadly yellow at sides. Genital valves darkened, surpassing tuberculum anale and cerci for about half length of segment 10, yellow-tipped; cerci pricker-shaped, yellow. Measurements: abdomen (incl. valves) 35.5, hind wing 25.0, pterostigma 1.0 mm. Female (juvenile, red colour phase). Ground colour of whole body bright orange- pink; dark markings much as in male, differing as follows. Labrum wholly pale save for a deep TIJDSCHRIET VOOR ENTOMOLOGIE, DEEL 129, AFL. 9, 1986 (1987) black midbasal spot; front of face likewise, ex- cept postclypeus deep black. Dorsum of pro- thorax with a fine X-shaped black mark and a median spot at extreme base of its posterior lobe, the latter shaped much as in the adult, but | side-lobes more rounded. Synthorax with the metallic green dorsal band a little narrower, standing out clearly on either side of the median carina, which remains orange, as also the ante- alar triangles. Abdomen as in the adult female, but end segments 9—10 for the greater part orange. [ REFERENCES | Brauer, F., 1868. Dritter Bericht über die von Herrn G. Semper mitgeteilten, von dessen Bruder auf den Philippinen gesammelten Neuropteren und Beschreibung einer neuen Libellen-Gattung aus dem Museum Godeffroy in Hamburg. — Ver- handlungen der K. K. zoologisch-botanischen Ge- sellschaft in Wien 18: 541—558. Campion, H., 1924. Descriptions of two new species of Teinobasis (Odonata). — Annals and Magazine of Natural History 9 (14): 612—614. Davies, D. A. L., & P. Tobin, 1984. The dragonflies of the world: A systematic list of the extant spe- cies of Odonata. Volume 1. Zygoptera, Anisozy- goptera. — Rapid Communications Societas In- ternationalis Odonatologica, Suppl. 3: i—ix, 1— 127. Kimmins, D. E., 1934. A note on two species of Tez- nobasis (Odonata) described by the late Herbert Campion. — The Entomologist 67: 197. Kimmins, D. E., 1957. Odonata collected by Mr. J. D. Bradley on Guadalcanal Island, 1953-54. — Bul- letin of the British Museum (Natural History), se- ries Entomology 5: 309-320. Kimmins, D. E., 1970. A list of the type-specimens of Odonata in the British Museum (Natural Histo- ry). Part III. — Bulletin of the British Museum (Natural History), series Entomology 24: 173— 205. Lieftinck, M. A., 1932. The dragonflies (Odonata) of New Guinea and neighbouring islands. I. Descrip- tions of new genera and species of the families Lestidae and Agrionidae. — Nova Guinea 15: 485-602. Lieftinck, M. A., 1935. The dragonflies (Odonata) of New Guinea and neighbouring islands. III. De- scriptions of new and little known species of the families Megapodagrionidae, Agrionidae and Li- bellulidae (genera Podopteryx, Argiolestes, Papua- grion, Teinobasis, Huonia, Synthemis and Procor- dulia). — Nova Guinea 17 (Zool.): 203—300. Lieftinck, M. A., 1937. The dragonflies (Odonata) of New Guinea and neighbouring islands. IV. De- scriptions of new and little known species of the families Agrionidae (sens. lat.), Libellulidae and Aeshnidae (genera /diocnemis, Notoneura, Papua- Liertinck: New Guinea and Solomon Islands Odonata 291 grion, Teinobasis, Aciagrion, Bironides, Agyrta- cantha, Plattycantha, and Oraeschna). — Nova Guinea (N. S.) 1: 1—82. Lieftinck, M. A., 1938. The dragonflies (Odonata) of New Guinea and neighbouring islands. V. De- scriptions of new and little known species of the families Libellaginidae, | Megapodagrionidae, Agrionidae (sens. lat.), and Libellulidae (genera Rbinocypha, Argiolestes, Drepanosticta, Notoneu- ra, Palaiargia, Papuargia, Papuargion, Teinobasis, Nannophlebia, Synthemis, and Anacordulia). — Nova Guinea (N.S.) 2: 47—128. Lieftinck, M. A., 1949. Synopsis of the odonate fauna of the Bismarck Archipelago and the Solomon Is- lands. — Treubia 20: 319—374. Lieftinck, M. A., 1953. Additions to the odonate fau- na of the Indo-Australian Archipelago. — Treubia 22: 233—269. Lieftinck, M. A., 1957. On some old types of coen- agrionine Odonata described from the Philippine Islands, with notes on allied species. — Zoolog- ische Mededelingen, Leiden 35: 161—175. Lieftinck, M. A., 1962. Odonata. — Insects of Micro- nesia 5 (1): 1—95. Lieftinck, M. A., 1963. Contributions to the odonate fauna of the Solomon Islands, with notes on zy- gopterous larvae. — Nova Guinea (Zool.) 21: 523—542, 1 plate. Needham, J. G., & M. K. Gyger, 1939. The Odonata of the Philippines, II. Suborder Zygoptera. — The Philippine Journal of Science 70: 239—314, plates 11—22. Selys Longchamps, E. de, 1877. Synopsis des Agrio- nines. 5me légion: Agrion (suite et fin). — Bulletin de l’Académie Royale de Belgique serie 2, 43 (2): 97—159 (separate p. 1—65). “wae dut ÿ o. sun DEN toe! Sti ee È = - x une È A + È NN = es agr st A | gia Sake! vange mat gia M i ; 5 sie ane ‘dice N aa | 3 es Ga tb heden ping > i ri Als ee shana Dieta RISTRETTO SIE pec! rout pe pri whois a pairks berk Mackie ach ee cha ai BE re Lbr ean plee a TIJDSCHRIFT VOOR ENTOMOLOGIE UITGEGEVEN DOOR DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING REGISTER VAN DEEL 129 * Een sterretje duidt een naam aan die nieuw is voor de wetenschap; cursieve paginanummers verwijzen naar de beschrijvingen van nieuwe taxa. * An asterisk denotes a name new to science; numbers in italics refer to descriptions of new taxa. ACARI aesculi, Neophyllobius 191 agrifoliae, Neophyllobius 191, superbus, Tycherobius 206-208 texanus, Ixodiphagus 181-183 theilerae, Ixodiphagus 181 [201 *Tillandsobius 191, 194, 195, 205 australis, Camerobia 191, 192, [195, 196, 198, 199 Camerobia 191, 192, 194, 195, [201 citri, Neophyllobius 191, 201 *Decaphyllobius 191, 192, 194, [195, 201 elegans, Neophyllobius 214 floridensis, Neophyllobius 191, [205 floridensis, Tillandsobius 207, [209 *gersoni, Decaphyllobius 191, 192, [200, 201, 202-204 hadros, Neophyllobius 191, 201, [202 hirtus, Ixodiphagus 183 hookeri, Ixodiphagus 181, 182 Hunterellus 181, 183 Ixodiphagus 181, 183 lombardinii, Neophyllobius 191, *Tycherobius 191, 194, 195, 205, aphrodite, Tipula 7 appendiculata, Nephrotoma 2 argyriventris, Bequaertiana, 23, [25 ariadne, Tipula 7 [208, 214 artemis, Tipula 7 virginiensis, Neophyllobius 191, asetosa, Melanophora 24, 32 [208 aspromontensis, Tipula 4 virginiensis, Tycherobius 205, aterrima, Oplissa 23, 24, 30, 31 [206, 208, 213 atramentaria, Stevenia 17, 19, 23, ARANEAE cornutus, Araneus 17 COLEOPTERA Anthrenus 243 carcharias, Saperda 17 Cetonia 77,98, 104 Geotrupes 98, 104 violaceum, Callidium 17 DICTYOPTERA nigra, Ostrava 107, 109 wolfforum, Mazonopterum 107 [205, 206 DIPTERA lombardinii, Tycherobius 206- [208, 210 *monspeliensis, Camerobia 191, [195, 199, 201 Neophyllobius 191-195, 214 Obdulia 201 *pistaciae, Camerobia 191, 195, [ 196, 198, 199 *polonicus, Tycherobius 191, 206, [207, 208, 212 rhytis, Neophyllobius 191, 205, [214 rhytis, Tycherobius 206 southcotti, Camerobia 191, 195, [198, 201 *stramenticola, Tycherobius 191, [206, 207, 208, 211 superbus, Neophyllobius 191, [206, 208 acerba, Angioneura 20, 21 Acompomintho 23, 27, 29, 30 aculeata, Nephrotoma 2 acuminata, Tipula 11 Acutipula 2, 6 alpina, Tipula 4 alpium, Tipula 3 analis, Nephrotoma 2 Angioneura 15, 20, 21, 23, 27 angustifrons, Phyto 17, 23, 18, [25, 26 angustifrons, Stevenia 23, angustifrons, Tromodesia 23, 25, [26 anicilla, Tipula 6 animula, Tipula 6 Anoplisa 30 anthe, Tipula 7 antichasia, Tipula 6 [31 atrata, Tanyptera 2 austriaca, Tipula 4 Azaisia 23, 27, 29, 30, 32 balcanica, Tipula maxima 2, 5 Baniassa 18, 20, 23, 25-27, 32 basilewskyi, Bequaertiana 23 benesignata, Tipula 4 Bengalia 16, 22 Bequaertiana 17, 23, 24, 26, 32 Beringotipula 2 betulae, Semudobia 125, 128, [132, 133, 136-140 Bezzimyia 20 bifasciculata, Tipula 7 bimacula, Tipula 5, 7, 9, 12 bimaculata, Dictenidia 2 bispina, Tipula 6 borystenica, Tipula 7 brevicornis, Paykullia 23 brevipalpis, Semudobia 126, 130, Ha, 135 brunneinervis, Tipula 5, 7, 11 bulbosa, Tipula 7 caesia, Tipula 3 Callidesia 23, 25, 26 Calliphora 77, 98 Calobataemyia 22 capensis, Melanomyoides 23, [27-29, 32 capensis, Rhinomorinia 23 capreola, Tipula 6 carbonaria, Metoplisa 23, 30-32 carpatica, Tipula excisa 4 294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, 1986 (1987) | cassiope, Tipula 6 fimbriata, Angioneura 21 caudatula, Tipula 6 festiva, Ctenophora 2 caudispina caudispina, Tipula 6 flaveolata, Ctenophora 2 caudispina parnonensis, Tipula 6 flavescens, Nephrotoma 2 Chaetostevenia 27 flavipalpis, Nephrotoma 2 cheethami, Tipula 4 flavipes, Queximyia 23, 28, 32 ciconia, Tipula truncata 6 flavolineata, Tipula 2 cilicrus, Clinodiplosis, 125 fulvipennis, Tipula 2 cinerascens, Tipula 6 furcula, Tipula 7 cinerella, Tipula 7 fuscicosta, Tipula 7 cingulata, Phyto 23 fuscipes, Dolichopeza 4 circumdata, Tipula 4 gimmerthali, Tipula 4 Clans, 175 22, 23, 27 glacialis, Tipula 4 Clinodiplosis 126 goriziensis, Tipula 4 clio, Tipula 6, 10 graeca, Dolichopeza 4 coerulescens, Tipula graecolivida, Tipula 7 latemarginata 3 griseithorax, Comoromyia 23, 30, Comoromyia 23, 30 [32 congolensis, Parazamimus 23, 26, grisescens, Tipula 4 [32 guestfalica, Nephrotoma 2 conspurcata, Helicella 17 guttata, Ctenophora 2 cornicina, Nephrotoma 2 harmonia, Tipula 6 couckei, Tipula 3 helicivora, Melanophora 17 crassiventris, Tipula 4 helvola, Tipula 2 cretis, Tipula 5, 7 hemiptera, Tipula strobliana 4 crocata, Nephrotoma 2 hera, Tipula 6 croceiventris, Nephrotoma 2 heros, Tipula 7 Ctenophora 2 hertingi, Phyto 27 cyrtoneurina, Angioneura 20,21 hirtigena, Stevenia 23, 30 danieli, Tipula 6 Hoplisa 30 Dasineura 125, 126, 136, 139 hortorum, Tipula 3 deceptoria, Stevenia 23 invenusta subinvenusta, Tipula 4 decipiens, Tipula 3 interbracta, Dasineura 130, 136, decolor, Tipula 7 [137 Dendrotipula 2 interserta, Tipula 4 Dictenidia 2 irrorata, Tipula 3 discrepans, Phyto 17, 21, 23 ismene, Tipula 6 Dolichopeza 4 istriana, Tipula 7 dorsalis, Nephrotoma 2 italica errans, Tipula 5, 7, 9 elegans, Ctenophora 2 jeekeli, Tipula 4 eleonorae, Tipula 4 jordansi, Tipula pannonia 7 Emodotipula 4 klytaimnestra, Tipula 6 engeli, Tipula 5, 7 kugleri, Paykullia 23 erato, Tipula 7 laetabilis, Tipula 4 errans, Tipula italica 5, 7, 9 lanispina, Tipula 6 euchroma, Nephrotoma 6 latemarginata coerulescens, eugeniana, Tipula 6 Tipula 3 eurykleia, Tipula penelope 6 lateralis, Tipula 3 euterpe, Tipula 6 latifurca, Tipula transcaucasica 6 excisa carpatica, Tipula 4 leandros, Tipula 7, 10 excisa excisa, Tipula 4 leda, Tipula 6 fascicula, Tipula 6 lepida, Rhinophora 17, 23, 26, fascingulata, Tipula 4, 10 [28, 32 fascipennis, Baniassa 23, 25, 27 Lestodiplosis 126 fascipennis, Tipula 2 leto, Tipula 6 fastidiosa, Dasineura 136, 137 limbata, Tipula 4 fastuosa, Ctenophora 2, 3 limitata, Tipula 4 fernandezi, Stevenia 23 livida, Tipula 2, 11 lobata, Acompomintho 23, 29, 32 longimana, Macrotarsina 23, 29, [32 luna, Tipula 2 lunata, Tipula 2, 11 Lunatipula 2, 4-6, 8-12 lunulicornis, Nephrotoma 2 luridorostris, Tipula 4 luteipennis, Tipula 3 macropeliostigma, Tipula 7 macroselene macroselene, Tipula iy, macroselene pan, Tipula 7 Macrotarsina 23, 29, 40 maculata, Paykullia 17, 23, 25 marginella, Tipula 3 marshalli, Termitoloemus 22 maxima balcanica, Tipula 2, 5 mayerduerii, Tipula 4 Mediotipula 4 mediterranea, Tipula 5, 7 melanocephala, Phyto 17, 23, 26 Melanonya 15, 21, 22 Melanomyoides 23, 27, 29 Melanophora 17, 18, 20, 23, 24 melanoptera, Morinia 21, 22 mellea, Tipula 2 melpomene, Tipula 6 Metopisena 23 Metoplisa 23, 29, 30 milichioides, Ventrops 23, 27, 28 monstrabilis, Tipula 7 montana, Tipula 4 montium, Tipula 3 Morinia 15, 22 nana, Melanomya 21 nausicaa, Tipula 7 Nephrotoma 2, 4, 6 neurotica, Tipula 4 nigerrima, Morinia 22 nigra, Nigrotipula 2 nigricornis, Tanyptera 2 Nigrotipula 2 nodicornis, Tipula 3 nubeculosa, Tipula 3 Nyctia 22 obscura, Angioneura 20, 21 obscura, Azaisia 23, 29 obsoleta, Tipula 3 Odonatisca 3 oldenbergi, Oplisa 30 oleracea, Tipula 3 onusta, Tipula 7 Oplisa 23, 29, 30, 32 Opsodexia 20 orientalis, Tipula 5, 7 ornata, Ctenophora 2 pabulina, Tipula 3 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, 1986 (1987) pachyprocta, Tipula 7 pallidicosta, Tipula 4 paludosa, Tipula 3 pan, Tipula macroselene 7 pandorra, Tipula 7 pannonia jordansi, Tipula 7 pannonia pannonia, Tipula 7 Parazamimus 20, 23, 24, 26, 27 parnonensis, Tipula caudispina 6 paucipila, Baniassa 23, 26 pauciseta, Phyto 23, 25 Paykullia, 17-20, 23, 24, 26, 27, [32 pectinicornis, Ctenophora 2 pelidne, Tipula 7 peliostigma, Tipula 3, 11 penelope eurykleia, Tipula 6 penelope penelope, Tipula 6 Phyto 15, 17-20, 22, 23, 26, 27, [30, 32 pictipennis, Callidesia 23, 25, 32 pierrei, Tipula 3 pinnifer, Tipula 6 Platytipula 3 plitviciensis, Tipula 4 pollinosa, Oplisa 23, 30 polydeukes, Tipula 6 praecox, Tipula 6 pratensis, Nephrotoma 2 profdrassi, Tipula 6 pruinosa, Tipula 3 pseudoirrorata, Tipula 4 pseudopruinosa, Tipula 4 pseudovariipennis, Tipula 3 Pterelachisus 3, 4 pythia, Tipula 6 quadrifaria, Nephrotoma 2 quadristriata, Nephrotoma 2 Queximyia 23, 27, 29 Rhinomorinia 22, 23, 26, 27, 29, [32 Rhinophora 17, 18, 23, 27, 29 riedeli, Tipula 7 roralis, Melanophora 17, 21, [ 23-25, 32 rubricosa, Tricogena 17, 23, 31, [32 rufina, Tipula 3 rufula, Tipula 6 saginata, Tipula 4, 5 sarajevensis, Tipula 4 Sarcophaga 19 sarcophagina, Rhinomorinia 17, [23, 26, 28 Savtshenkia 3, 4 savtshenkoi, Tipula 7 scalaris, Nephrotoma 2 schmidti, Tipula 6 Schummelia 3, 4 scripta, Tipula 3 scurra, Nephrotoma 2 seguyi, Tipula 6 selene, Tipula 3 Semudobia 125, 132, 133, 135, [ 136, 139, 140 setitarsis, Azaisia 23 Shannoniella 20 siebkei, Tipula 4 sigma, Tipula 6 simulans, Tipula 4 Siphona 16 skuhravae, Semudobia 126, 128, [ 132, 136-140 soosi, Tipula 5, 7, 9, 12 Stevenia 17, 18, 22, 23, 26, 27, [29, 30, 32 stigmatella, Tipula 4 strobliana hemiptera, Tipula 4 strobliana strobliana, Tipula 4 stubbsi, Tipula 3, 5 subcunctans, Tipula 3 subinvenusta, Tipula invenusta 4 submaculosa, Nephrotoma 2 submarmorata, Tipula 3 submontium, Tipula 3 subnodicornis, Tipula 4 subsignata, Tipula 4 subtrunca, Tipula 6 subtruncata, Tipula 6 Tanyptera 2 tarda, Semudobia 126, 128, 132, [133, 136-140 tenuicornis, Tipula 2 tenuipes, Nephrotoma 4 tergestina, Oplisa 23, 30 Termitoloemus, 15, 22 thais, Tipula 6 theia, Tipula 6 tibonella, Tipula 7 Tipula 2-4 titania, Tipula 6 transcaucasica latifurca, Tipula 6 Tricogena 17, 18, 23, 26, 29, 30 Tromodesia 23, 25-27, 32 trunca, Tipula 6 truncata ciconia, Tipula 6 truncata truncata, Tipula 6, 10 truncorum, Tipula 3 tyche, Tipula 6 umbratica, Stevenia 17, 23 unca, Tipula 2 urania, Tipula 7 variicornis, Tipula 3 varipennis, Tipula 3 Ventrops, 23, 27, 32 vernalis, Tipula 3, 5 295 Vestiplex 3, 4 vibripennis, Tromodesia 25 vittata, Tipula 2 vogtenhuberi, Tipula 7 vorax, Lestodiplosis 125 wewalkai, Tipula 7 wolfi, Tipula 7 xanthocephla, Rhinomorinia 23, [28 Yamatotipula 3, 7 zonaria, Tipula 4 EPHEMEROPTERA Caenis 88 Ephemera 81, 87 Oligoneuriella 88 Ostrava 109 HOMOPTERA *brendelli, Lembeja 141, 142, 146, [147, 176, 178-180 *dekkeri, Lembeja 141, 142, [ 145-147, 150, 155-158, 173 *distanti, Lembeja 141, 142, 145- [147, 170, 173, 174, 175, 178, 180 *elongata, Lembeja 141, 142, 144, [146, 147, 150, 153-155, 157, [158, 173 faticina, Prasia 141 fatiloqua, Lembeja 141, 142 foliata, Cephaloxys 141, 149, 150 foliata, Lembeja 141-147, [ 149-151, 153, 158, 160, 162, PGO ONS WA foliata, Prasia 149 fruhstorferi, Lembeja 141, 142, [158 harderi, Lembeja 141, 149 *hollowayi, Lembeja 141, 142, 146, 148, 162-164, 166, 173, 177 *incisa, Lembeja 141, 142, 146, 148, 158, 166, 170, 171-173 Iruana 142 Lembeja 141, 142, 158 maculosa, Lembeja 141, 142, 160 maculosa, Perissoneura 178 *majuscula, Lembeja 141, 142, [144, 146, 148, 169, 170, (171, 173, 179 *minahassae, Lembeja 141, 142, [ 145-147, 150, 151-153, ISS} 775) 1177 *mirandae, Lembeja, 141, 142, [145, 146, 148, 166-168, 170-173 *oligorhanta, Lembeja 141, 142, [146, 148, 158, 164-166, 173 papuensis, Lembeja 141, 142 paradoxa, Lembeja 141, 142 296 *pectinulata, Lembeja 141, 142, [144, 146, 148, 158-160, 162, 173 Perissoneura 141 Prasia 141, 150 robusta, Lembeja 141, 142 roehli, Lembeja 141, 142 *sangihensis, Lembeja 141, 142, [146, 148, 149, 160-162 sanguinolenta, Lembeja 141, 142, [178 tincta, Lembeja, 142 tincta, Prasia, 142 ~ typica, Jacatra 143, 160 vitticollis, Lembeja 142 vitticollis, Prasia 141 HYMENOPTERA affinis, Pelopoeus 254 affinis, Sphex 254 Apis 77 atripes, Pelopoeus 240 atripes, Sceliphron deforme 229, [233, 234, 238, 240, 242, 253 betulae, Misocyclops 125 betulae, Platygaster 125-128, [ 132-134, 136, 137, 139, 140 betularia, Platygaster 125-128, [ 131-134, 136, 137, 139, 140 bicinctum, Sceliphron deforme [249 bicinctum, Sceliphron rufopictum [244, 249, 254 bilineatum, Sceliphron 228 bruijnii, Sceliphron 256 bruinjnii, Pelopoeus 256 bruinjnii, Sceliphron 256 bruinjnii, Sceliphron formosum [251, 254, 256, 257, 259 bruynii, Sceliphron 256 Chalybion 219, 221 coromandelicus, Pelopoeus 243 coromandelicum, Sceliphron [218-220, 222, 224, 239, 243, 252 curvatus, Pelopoeus 228, 230 curvatum, Sceliphron 218-221, [224, 227, 228, 230, 234, [238, 252, 256 deforme, Sceliphron 218-221, [224, 229-231, 234, 236, [238, 240, 242, 243, 253 deforme atripes, Sceliphron 229, [233, 234, 238, 240, 242, 253 deforme bicinctum, Sceliphron [249 deforme deforme, Sceliphron [231, 232, 234, 236, [238, 240, 249, 253 *deforme femorale, Sceliphron [217, 229, 231, 234, [236, 240, 243, 253 deforme japonicum, Sceliphron [240 deforme koreanum, Sceliphron [240 deforme nippo nicum, Sceliphron 233, 234, 240, 253 deforme rufopictum, Sceliphron [249 deforme taiwanum, Sceliphron [238 deforme tibiale, Sceliphron 231, [234, 236, 238, 243, 253 deforme unifasciatum, Sceliphron 254 deformis, Pelopoeus 236 deformis, Sceliphron 252 fallax, Sceliphron 256 *femorale, Sceliphron deforme LA, 220) DIG 2345 [ 236, 240, 243, 253 fervens, Pelopoeus, 242 fervens, Sceliphron 219-221, 224, [230, 235, 237, 242, 243, 255 fervens murarium, Sceliphron [254 flavo-fasciatus, Pelopoeus 249, [251 formosum, Sceliphron 218-220, [224, 234, 243, 251, 256-258 formosum bruinjnii, Sceliphron BOL 2325 258, 2375 259 formosum formosum, Sceliphron [245, 251, 256-258 formosum koreanum, Sceliphron [240 formosum ocellare, Sceliphron [248, 251, 256-258 formosus, Pelopoeus 258 formosus, Sceliphron 258 funestum, Sceliphron 219, 220, (223-226, 228, 243, 251 hiemalis, Platygaster 136 japonicum, Sceliphron deforme [240 *kalshoveni, Sceliphron rufopictum 217, 219, 220, 224, [241, 244, 249, 255 koreanum, Sceliphron deforme [240 koreanum, Sceliphron formosum [240 *laticinctum, Sceliphron rufopictum 217, 218, 241, 245, 248, 249, 252, 253, 255 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, 1986 (1987) lineatipes, Sceliphron 234, 236 mamertes, Piestopleura 126 Metaclisis 129, 130, 132, 135, [138, 140 Misocyclops 125 mocsaryi, Sceliphron 254 murarium, Sceliphron 219, 220, [224, 246, 254, 255 murarium, Sceliphron fervens [254 murarius, Pelopoeus 254 nipponicum, Sceliphron deforme [233, 234, 240, 253 ocellare, Sceliphron 258 ocellare, Sceliphron formosum [248, 251, 256-258 -papuanum, Sceliphron 258 Pelopoeus 224, 258 phragmitis, Metaclisis 125-127, [130, 131, 133, 136, 137, 139, 140 Piestopleura 129, 136 Platygaster 125, 127-130, 132, [133, 135-140 Prosceliphron 217-222, 224, 225, [228, 230, 235, 238, 240, 242, [243, 249, 254, 256, 258 pulchellum, Sceliphron 228 pulchellum, Sceliphron rectum [228, 251 rectum, Sceliphron 218-220, [ 223-226, 228, 243 rectum pulchellum, Sceliphron [228, 251 rectum rectum, Sceliphron 228, [251 rufipes, Pelopoeus 254 rufipes, Sceliphron 254 rufopictum, Sceliphron 241, 243, [244, 247, 249, 254, 255 rufipictum, Sceliphron deforme [249 rufopictum bicinctum, Sceliphron [244, 249, 254 *rufopictum kalshoveni, Sceliphron 217, 219, 220, 224, [241, 244, 249, 255 *rufopictum laticinctum Sceliphron 217, 218, 241, 245, [248, 249, 252, 253, 255 rufopictum rufopictum, Sceliphron 241, 249, 253-255 rufopictus, Pelopoeus 249 rufopictus 238 Sceliphron 217-221, 226 shestakovi, Sceliphron 219-224, [251 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 129, 1986 (1987) 297 taiwanum, Sceliphron deforme Gomphus 115 [238 grandis, Aeshna 109 tibiale, Sceliphron 234, 236 gurneyi, Papuagrion 263, 278, tibiale, Sceliphron deforme 231, [280 [234, 236, 238, 243, 253 gurneyi, Teinobasis 279, 281, unifasciatum, Sceliphron 219, [282, 287 [220, 224, 244, 246, 254, 255 Hemianax 64 unifasciatum, Sceliphron Ictinogomphus 115 deforme 254 Idiocnemis 266 unifasciatus, Pelopoeus 254 *imitans, Teinobasis 263, 265, [ 278-282, 286, 288 LEPIDOPTERA imperator, Anax 62-64, 66, 68, farinalis, Pyralis 17 [69, 72, 103, 109 neustria, Malacosoma 17 inornata, Idiocnemis, 275 tabaniformis, Paranthrene 17 ‘*isabellae, Lieftinckia 263, 265, [273, 274, 279 ODONATA juncea, Aeshna 98 Aeshna 68 kimminsi, Lieftinckia 265, 274, aluensis, Teinobasis 263, 265, 12755 2776 278 [278-281, 286-288 lairdi, Lieftinckia 265, 271-275, Amphicnemis 280 [278 Anax 61, 64 Lieftinckia, 265, 268, 271, 273, Archibasis 280 [279 bradleyi, Teinobasis 265, 278, malacodora, Lochmaeocnemis [283-285, 288 [264 cancellatum, Orthetrum 103 *malaitae, Lieftinckia 263, 265, *chionopleura, Teinobasis 263, (272,275 [265, 278, 283-285, 286 mirabilis, Hemiphlebia 110 Coeliccia 274 mixta, Aeshna 36 cyanea, Aeshna 36, 59, 64, 70, 97, | Nesobasis 280 [100, 103,115 obliterata, Idiocnemis 264 Davidius 115 *obtusilingua, Teinobasis 263, emarginata, Teinobasis 263, [265, 278, 283, 284, 288 [279-282 Oligoneuriella 88 ephippiger, Hemianax 64 Onychogomphus 115 Epiophlebia, 36, 39, 42, 44, 56, Papuagrion 278-280 [61, 101, 109, 115, 118, 119 papuensis, Hemianax 64 erythrostigma, Paramecocnemis Paramecocnemis 266 [264 Pericnemis 280 filicornis, Torrenticnemis 264 Procordulia 263 frequens, Sympetrum 92 *prothoracica, Rhyacocnemis 263, *gerdae, Salomocnemis 263, 265, [265-267 [269,277 puella, Coenagrion 64 Pseudagrion 280 *ramosa, Lieftinckia 263, 265, 274, [275, 276, 279 Rhyacocnemis 265 rufithorax, Teinobasis 278 *Salomocnemis 263, 265, 268 salomonis, Lieftinckia 265, 268, [269, 271, 272, 277, 278 Seychellibasis 278 *simulans, Teinobasis 263, 265, [ 278-282, 286, 287 stictica, Pericnemis 280 striolatum, Sympetrum 101 sufficieus, Rhyacocnemis 265-268 superstes, Epiophlebia 117 Teinobasis 263, 265, 278-281, [284, 286, 287 Torrenticnemis 263 ORTHOPTERA Locusta 77, 97, 98, 104 Schistocerca 92, 104, 114 MOLLUSCA elegans, Succinea 21 PLANTAE Betula 125, 135 Betula populifolia 135 Casuarina 275, 283 Corylus avellana 198 Cyclops 202 Eucalyptus 195 Eucalyptus camaldulensis 195 Hedera himalaica 214 Pistacia 198, 199 Platanus acerifolia 201 Quercus agrifolia 202 Salvadora oleoides 202 Tamarix aphylla 201 a Jap fea Serbe 4 si durati 0 ERI CE ud Hist nate ol I ee ; Li x en eine? > = Pi Si 5 2, in ss a : e. 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