Research Article |
Corresponding author: Xingyue Liu ( xingyue_liu@yahoo.com ) Academic editor: Susanne Randolf
© 2015 Ulrike Aspöck, Xingyue Liu, Horst Aspöck.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Aspöck U, Liu X, Aspöck H (2015) The Dilaridae of the Balkan Peninsula and of Anatolia (Insecta, Neuropterida, Neuroptera). Deutsche Entomologische Zeitschrift 62(2): 123-135. https://doi.org/10.3897/dez.62.5199
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Basing upon all available information on type material of Dilar turcicus Hagen, 1858, Dilar syriacus Navás, 1909, and Dilar lineolatus Navás, 1909, together with a large number of dilarid specimens, the pleasing lacewings of Anatolia and Southeast Europe are revised. The current taxonomic concept of D. turcicus is confirmed, and a lectotype is designated. Dilar turcicus is widely distributed in the southeast of Europe (being the only representative of Dilaridae in this region), in Anatolia, and, most probably, in the Caucasus region. Dilar syriacus and D. lineolatus remain nomina dubia. Dilar syriacus might occur in Anatolia, while D. lineolatus is a species occurring in western Central Asia. Two new species, Dilar anatolicus sp. n. and Dilar fuscus sp. n. are described from Anatolia. Wings and genital segments of the three species occurring in Anatolia are illustrated, and a map documenting the known distribution of these species is provided.
Dilar , lectotype designation, new species, nomina dubia, Europe, Asia minor
The Dilaridae is a small family of the order Neuroptera comprising about 100 described valid species worldwide. Most species – all assigned to the genus Dilar Rambur, 1838 – have been recorded in the Northern Hemisphere, and particularly in Central, East and Southeast Asia (
In the Western Palaearctic most species of Dilar have been found in the southwest of Europe in the Iberian Peninsula (
For many years we have, however, been aware of the existence of at least two further Dilar species which had been collected by two of us (H.A., U.A.) in Anatolia. However, the fact that the type material of D. turcicus had never been examined and the possibility of the occurrence of other described but not identifiable species (nomina dubia) prevented us from describing our unidentified species.
In the course of recent studies on Dilaridae of Asia, we have again examined the situation and have tried to clarify all open questions as far as possible, so that we can now present a summarising overview on the Dilaridae of Anatolia and of Southeast Europe.
Specimens (partly pinned and dried, partly preserved in alcohol) on which this study is based are deposited in the following collections: Collection of Horst & Ulrike Aspöck, Vienna, Austria (HUAC); Collection of Hubert & Renate Rausch, Scheibbs, Austria (HRRC); Natural History Museum Vienna, Austria (NHMW); Museum für Naturkunde der Humboldt-Universität zu Berlin, Germany (MFN); Muséum National d’Histoire Naturelle, Paris, France (MNHN); Muséum d’Histoire Naturelle, Genève, Switzerland (MHN); Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. (MCZ).
Genitalic preparations were made by clearing the apex of the abdomen in a cold, saturated KOH solution for 3-4 h. After rinsing the KOH with acetic acid and water, the apex of the abdomen was transferred to glycerine for further dissection and examination. Habitus photos were taken by using Nikon D60 and D800 digital camera with Nikon MICRO NIKKOR 105 mm lens, and the genitalic figures of the new species were made by hand drawing under Leica M165C stereo microscope. The genitalia of the lectotype of D. turcicus were photographed with a Leica DFC camera attached to a Leica MZ16 binocular microscope and processed with the help of Leica Application Suite. They were then stacked with ZereneStacker 64-bit and processed with Adobe Photoshop Elements 8. The terminology of the genitalia generally follows
Distribution map: Localities were taken from original literature (Suppl. material
Dilar Rambur, 1838: 9. Type species: Dilar nevadensis Rambur, 1838: pl. 9 (monotypy).
Cladocera Hagen, 1860: 56. Nomen nudum.
Lidar Navás, 1909: 153. Type species: Dilar meridionalis Hagen, 1866: 295, original designation.
Fuentenus Navás, 1909: 154. Type species: Dilar campestris Navás, 1903: 380, original designation.
Nepal Navás, 1909: 661. Type species: Nepal harmandi Navás, 1909: 661, original designation.
Rexavius Navás, 1909: 664. Type species: Dilar nietneri Hagen, 1858: 482, subsequent designation by Navás, 1914: 10.
A description of the genus has been given on several occasions in recent publications (
Dilar turcicus Hagen, 1858:
Dilar corcyraeus Navás, 1909:
After the original description by
D. turcicus shows a considerable variation in size and coloration of wings. Lengths of forewings vary from 7.5 to 13.5 mm. A reliable identification seems – at least presently – only possible on the basis of the male genital segments.
As far as we know the southeast of Europe does not harbour any other species besides D. turcicus. We have examined more than 200 specimens from various localities on the Balkan Peninsula and have not found any hint for the occurrence of a second Dilar species. Thus, it is a reasonable assumption that all published records of D. turcicus from localities situated in Europe really refer to this species. We have therefore included all published European records of D. turcicus in the map (Fig.
The exact type locality of D. turcicus remains unknown. In the middle of the 19th century the name Armenia was used also for regions in the east of Anatolia. Whether D. turcicus occurs in Armenia of present time is not sure. However,
Dilar corcyraeus has already been synonymised 35 years ago (
The biology of D. turcicus is poorly known, with only a few scattered observation records (
Dilar syriacus Navás, 1909:
This species has been described based on a single male specimen without abdomen deposited in the MNHN, Paris. According to the description of
Dilar lineolatus Navás, 1909:
This species was described on the basis of a specimen from “Turkmenien (Tekke)”. Turkmenien is the old German word for that part of western Central Asia which is largely identical with the present Turkmenistan. In his description of the species
The holotype of D. lineolatus (Figs
This species is characterized by the forewings with numerous pale brown spots and by the male gonocoxite complex 9, 10 and 11 with short, feebly sclerotized ninth gonocoxite and elongate, blade-like tenth gonocoxite.
Male. Length of forewing 11.0–12.2 mm, of hindwing 9.5–10.3 mm.
Head pale yellowish brown, with pale yellow setose tubercles. Compound eyes blackish brown. Antenna with ca. 29 segments, pale yellow, with scape and pedicel pale brown, flagellum unipectinate on most flagellomeres, medial branches much longer than those on both ends, longest branch nearly 3.0 times as long as the corresponding flagellomere, but branch of 1st flagellomere short and dentate, distal eight flagellomeres simple.
Prothorax pale yellowish brown, pronotum dark brown with several hairy yellowish tubercles; meso- and metathorax yellow, each notum laterally with a pair of broad, brown markings and a pair of pale brown narrow stripes; mesonotum medially with a pair of additional brown markings near anterior margin. Legs pale yellow, with femora blackish brown at tip. Wings hyaline, slightly yellowish brown, with numerous pale brown spots. Forewing ~2.5 times as long as wide, densely spotted, with markings slightly darkened toward anal region; a few markings on posteroproximal area connected with each other, forming short transverse stripes; an immaculate area present distal to median nygma; two nygmata present on proximal portion, both slightly smaller than median nygma. Hindwing ~2.1 times as long as wide, slightly paler than forewing, with dark markings largely reduced; one nygma present at middle. Veins pale yellowish brown; Rs with four main branches on both fore- and hindwings.
Abdomen pale yellowish brown, pregenital segments dorsally brown. Ninth tergite in dorsal view with nearly truncate anterior margin and a deeply V-shaped posterior incision, leaving a narrow median portion and a pair of subtriangular hemitergites, which are obtuse and slightly incurved distally; in lateral view broad, with straight ventral margin and arcuate posterior margin. Ninth sternite obviously shorter than ninth tergite, with nearly truncate posterior margin. Ectoproct in dorsal view with a pair of digitiform projections, posteroventrally with a pair of subsemicircular and flattened lobes, a pair of bifid unguiform projections and a pair of short, feebly sclerotized, digitiform projections. Ninth gonocoxite short, shield-like, mostly membranous and transparent, with sclerotized lateral margin and with a strongly narrowed, obtuse apex; tenth gonocoxite slenderly elongate, blade-like, nearly 2.5 times as long as ninth gonocoxite, straightly directed posteriorly with median portion feebly curved; fused eleventh gonocoxites (= gonarcus) beam-shaped, laterally connectedto base of ninth gonocoxites. Hypandrium internum subtriangular, with lateral margins slightly arcuate.
Female. Length of forewing 10.9 mm, of hindwing 10.0 mm.
Seventh sternite in lateral view subtrapezoidal. Eighth abdominal segment without subgenitale. Basal part of bursa copulatrix sac-like, nearly rhomboid in ventral view, proximally membranous and strongly rugose, distally sclerotized into a pair of lateral sclerites and a median ridge. Ectoproct rather small, ovoid.
Holotype ♂, “Asia minor, Kizilçahamam [a district of Ankara Prov., 40°28′N, 32°38′E], 1200 m, 7. 70 / G. Friedel leg.” (HUAC). Paratypes: 1♀, same data as holotype (HUAC); 1♂, “ANATOLIEN, Icel, Kilik. Taurus, Namrun [a town of Toroslar Dist., Mersin Prov.], 37.03N, 34.46E, 1400-1800 m, 18.VI. / TÜRKEI, ANATOLIEN 1979, C. Holzschuh & F. Ressl” (HUAC); 1♂, “ANATOLIEN, Bursa, Uludağ, 700 m, 40.12N, 29.04E, 6.VII. / BULGARIEN-TÜRKEI-GRIECHENLAND-EXP. 1978, H. & U. ASPÖCK, H. & R. RAUSCH, P. RESSL” (HUAC); 3♂1♀, “As[ia]. Min[or]. Kizilcahamam 18-24.VII.[19]72. Pinker” (HRRC); 1♂, “ANATOLIEN, Icel, Kilik. Tauris, Namrun, 37.03N/34.46E, 1400-1800 m, 29.V.-3.VII.[19]79, TÜRKEI, ANATOLIEN 1979 C. Holzschuh, F. Ressl” (HRRC); 1♂, “Dilar syriacus ♂ Nav[ás], 620 Taurus, 48. Asie. Min[or]. Coll. Pictet” (MHN); 1♂, “Syrien, Ehrenberg/Lidar turcicus Hag/388/Type” [The word Type on the old label is wrong] (MFN).
Turkey: Western, central, and southern parts of Anatolia (Fig.
Adjective, masculine, nominative, singular; an attribute to the genus name. From lat. anatolicus 3 = referring or belonging to Anatolia, the Asian part of Turkey.
Dilar anatolicus sp. n. can be distinguished from all other western Palaearctic species of Dilar based on the slenderly elongate male ninth gonocoxites and the short, largely membranous male tenth gonocoxites. In appearance, D. anatolicus sp. n. looks similar to its co-existing Turkish species D. turcicus, but it can be easily distinguished from the latter by the male ninth tergite without median projection and by the different shape of sclerotization of the female bursa copulatrix. Interestingly, D. anatolicus sp. n. appears to be closely related to D. sinicus Nakahara, 1957, which is distributed in northern China (
Dilar anatolicus sp. n. 20. Male genitalia, dorsal view; 21. Male genitalia, ventral view; 22. Male genitalia, lateral view; 23. Male ectoproct, caudal view; 24. Female genitalia, lateral view; 25. Bursa copulatrix, ventral view. bc: bursa copulatrix; e: ectoproct; gx9–11: ninth to eleventh gonocoxites; hi: hypandrium internum; S7–9: seventh to ninth sternite; T7–9: seventh to ninth tergite. Scale bars = 0.5 mm.
This species is characterized by the entirely brown wings without distinct markings (at least in the female).
Female. Length of forewing 10.9–12.2 mm, of hindwing 9.9–11.2 mm.
Head brown, with three pale yellow setose tubercles; vertex medially with a blackish brown vittae. Compound eyes blackish brown. Antenna with ca. 25 segments, pale brown.
Prothorax blackish brown, pronotum with pale yellow anterior margin and several yellowish tubercles clothed with brownish hairs; mesothorax and metathorax blackish brown, dorsally yellow at middle. Legs yellowish brown, with each segment blackish brown at tip. Wings entirely brown, without distinct markings. Forewing ~2.5 times as long as wide, with distal half slightly paler; proximally with two or three nygmata, medially with one nygma, which is slightly larger than proximal ones. Hindwing slightly paler than forewing; one nygma present at middle. Veins brown; Rs with six main branches on both fore- and hindwings.
Abdomen pale brown. Ovipositor pale yellowish brown. Seventh sternite in lateral view subtriangular. Eighth abdominal segment without subgenitale. Basal part of bursa copulatrix sac-like, subtriangular in ventral view, most parts membranous and strongly rugose, with ventrolateral portions slightly sclerotized anteroposteriad. Ectoproct rather small, ovoid.
Male. Unknown.
Holotype ♀, “Prov. AYDIN, Str., Nazilli-Beydağ, 38.01N/28.18E, 650 m, 24.V./SUDWEST-ANATOLIEN-EXP, 1981, H. et U. et Ch. ASPÖCK, H. et R. RAUSCH, F. RESSL” (HUAC). Paratypes: 1♀, same data as holotype (HUAC); 1♀, “Prov. AYDIN, Str. Nazilli – Beydag, 38.01N/28.18E, 650 m, 24.V., SÜDWEST-ANATOLIEN-EXP. 1981, H. et U. et Ch. ASPÖCK, H. et R. RAUSCH, F. RESSL, 81/25” (HRRC); 1♀, “Dilar syriacus ♀ Nav[ás]. Cotypus, 620/48. Asie min[or]. Coll. Pictet” (MHN).
The only reliable record of D. fuscus sp. n. is from the type locality in western Anatolia (Fig.
Adjective, masculine, nominative, singular; an attribute to the genus name. From lat. fuscus 3 = dark. The name refers to the dark coloration of the wings.
Despite the unknown male, D. fuscus sp. n. is a spectacular new species and can be easily distinguished from all other species of Dilar based on the entirely brown wings without distinct markings. The female genitalia of D. fuscus sp. n. are principally similar to the co-existing species D. turcicus based on the basal part of bursa copulatrix with lateral portions slightly sclerotized. It can, of course, not be excluded that the hitherto unknown male has a different coloration of the wing membrane or even a patterned wing membrane. Such kind of sexual dimorphism has been known from other Dilaridae (
Distribution of Dilaridae in Southeast Europe and Anatolia (see geographical coordination and references of distribution records in Suppl. materials
Three species of the genus Dilar, i.e. D. turcicus, D. anatolicus sp. n. and D. fuscus sp. n., are now known from the Balkan Peninsula and Anatolia, with clear specific identities. In the southeast of Europe only D. turcicus occurs, while in Anatolia all these three species are present. Dilar fuscus sp. n. is known in the female only, which can easily be identified by the unusual dark coloration of the wings (see Figs
D. turcicus is a Pontomediterranean faunal element with a distribution comprising large parts of Southeast Europe, probably all major parts of Anatolia, and most probably parts of the Caucasus region. To the best of our knowledge D. turcicus is the only representative of the family Dilaridae in the Balkan Peninsula and in the southern parts of East Europe. The northernmost records of this species in Europe are in Croatia, Bulgaria, Ukraine, and Russian north of Caucasus. Most probably the species had survived during the last glacial period in various refugial areas in the southeast of Europe and in Anatolia respectively.
In Anatolia D. anatolicus sp. n. and D. fuscus sp. n. have been known to us for long, but with respect to the unsolved questions concerning the described species they remained undescribed until now. After examination, re-examination and re-consideration of the three pertinent species – Dilar turcicus Hagen, Dilar syriacus Navás, and Dilar lineolatus Navás respectively – the situation has largely been clarified (concerning D. syriacus and D. lineolatus as far as possible at present). Dilar syriacus (type locality: probably surroundings of Beirut) remains a nomen dubium, it cannot be excluded that it will turn out to be identical either with D. turcicus (then it would be a junior synonym) or with D. anatolicus sp. n. (then it would become a senior synonym). However, it may also be a species different from both.
D. lineolatus has erroneously been associated with Anatolia. The species has been described from the region of Tekke in Turkmenia (probably today in Turkmenistan) and does presumably not occur in Anatolia. The existing remains of the damaged holotype cannot be assigned to any of the known species of Dilar.
We want to express our grateful thanks to a number of colleagues who have helped substantially in this study: Philip D. Perkins and Rachel Hawkins (MCZ) organized, promoted and speeded the sending of photographs and finally the syntype of D. turcicus from Armenia which became the lectotype and thus the precondition for the taxonomy in this paper. Michael Ohl and Lukas Kirschey (MFN) have sent photographs of the other syntype of D. turcicus and of the type of D. lineolatus and a further specimen which turned out to represent D. anatolicus sp. n. Peter J. Schwendinger (MHN) provided specimens from the Pictet collection. Hubert and Renate Rausch (Scheibbs) entrusted their enormous Dilaridae collection to us for this study. Antoine Mantilleri (MNHN) helped us committedly to get into contact with the persons curating the Neuropterida collection. Several specimens of D. anatolicus sp. n. and of D. turcicus respectively in the HUAC were collected by the following entomologists: Ernst Arenberger, Georg Friedel †, Carolus Holzschuh, Friedrich Kasy †, Josef Klimesch †, Hans Malicky, Franz Ressl †, Alfred Radda and Josef Thurner †. Harald Bruckner (NHMW) deserves our special gratitude for providing the database of all the records, for producing the distribution map and finally for photographs, especially for the focus stacking images of the genital sclerites of the lectotypus of D. turcicus. We gratefully acknowledge the valuable input of the reviewers. This research was supported by the National Natural Science Foundation of China (Nos. 31322051 and 31320103902).
Hermann August Hagen (1817–1893) moved from Germany to Cambridge, Mass., USA in 1867, where he became the first professor of entomology in the USA. He took (at least parts of) his collection with him to the Museum of Comparative Zoology, where it is still preserved.
Sources of records upon which the distribution map is based
Data type: records data
Records on which the distribution map is based
Data type: records data