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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
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a,
ry FE x
1 Noe Banane >
==.
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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.
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|
|
|
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Fortsetzung. —
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DEEL 129 AFLEVERING 2 1986
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UITGEGEVEN DOOR
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| 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
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= 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.
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text. Genera recorded as woodlouse parasites are marked with an asterisk.
|
Pape: Rhinophoridae 33
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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
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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.
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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
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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
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MALES je
5
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e
10 * |
à |
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5 * |
ni |
*
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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
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FEMALES ®
15 e
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e
10 *
*
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5 * |
= |
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al |
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"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.
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