Revision of Chinese Dilaridae (Insecta: Neuroptera) (Part II): Species of the Genus Dilar Rambur from Tibet

Zootaxa 3878 (6): 551–562 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2014 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3878.6.3 http://zoobank.org/urn:lsid:zoobank.org:pub:7ADAE88A-0003-41FC-8795-8897001001B2 Revision of Chinese Dilaridae (Insecta: Neuroptera) (Part II): Species of the genus Dilar Rambur from Tibet

WEI ZHANG1, XINGYUE LIU1,5, HORST ASPÖCK2 & ULRIKE ASPÖCK3,4 1Department of Entomology, China Agricultural University, Beijing 100193, China. E-mail: [email protected] 2Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University of Vienna, Kinderspitalgasse 15, A-1095 Vienna, Austria. E-mail: [email protected] 3Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, A-1010 Vienna, Austria. E-mail: [email protected] 4Department of Integrative Zoology, University of Vienna, Althanstraße 14, A-1090 Vienna, Austria. E-mail: [email protected] 5Corresponding author. E-mail: [email protected]

Abstract

Three species of the genus Dilar Rambur, 1838, are recorded and described from Tibet in southwestern China, including Dilar geometroides H. Aspöck & U. Aspöck, 1968, Dilar harmandi (Navás, 1909), and Dilar tibetanus Yang, 1987, with the former two species recorded in Tibet for the first time. All three species are redescribed. Dilar aspersus Yang, 1988, and Dilar pusillus Yang, 1992, are synonymized with Dilar geometroides Aspöck & Aspöck, 1968, and Dilar tibetanus Yang, 1987, respectively. A key to the Dilar species from Tibet is provided.

Key words: Dilaridae, Dilar, Tibet, China

Introduction

Tibet, also known as the Xizang Autonomous Region, is a plateau region in Asia, northeast of the Himalayas in China and is commonly cited as the highest region on Earth, with an average elevation of 4,900 meters. (Yang 1987). The insect fauna in Tibet is particularly rich, but still poorly known. Remarkably, southeastern Tibet, containing tropical and monsoon rainforest, provides a variety of habitats to numerous Oriental insect species, a large number of which are endemic to this area (Huang & Han 1988; Yang et al. 2004). Hitherto, the lacewing family Dilaridae is known from Tibet with only three species, i.e. Dilar aspersus Yang, 1988, Dilar pusillus Yang, 1992, and Dilar tibetanus Yang, 1987 (Yang 1987, 1988, 1992; Oswald 2013). These species were described based on only external morphology but not genitalic characters, which are crucial for the specific identification. During our study of Dilaridae from Tibet, three species of the genus Dilar Rambur, 1838, were found. Based on the examination of the male genitalia, D. pusillus Yang was found to be conspecific with D. tibetanus Yang, while D. aspersus Yang is conspecific with D. geometroides Aspöck & Aspöck, 1968, which has previously been recorded from Nepal (Aspöck & Aspöck 1968). In addition, Dilar harmandi (Navás, 1909), which has previously been described from northeastern India (Navás 1909), is recorded from Tibet and Nepal for the first time. In the present paper, these three species of Dilar are re-described. In addition a key to the species of Dilar from Tibet is given. The present study on Dilaridae of Tibet is part of a revision of the family Dilaridae of China, which was started with a publication on the Dilaridae of northern China (Zhang et al. 2014).

Material and methods

Specimens for the present study are deposited in the Entomological Museum of China Agricultural University

Accepted by B. Price: 17 Oct. 2014; published: 30 Oct. 2014 551 (CAU), Beijing, Muséum National d’Histoire Naturelle (MNHN), Paris, France, the National Museum of Nature and Science (NSMT), Tokyo, Japan, Zoologische Staatssammlung München (ZSM), Munich, Germany, and the Collection of H. & U. Aspöck (HUAC), Vienna, Austria. 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 glycerin for further dissection and examination. Habitus photos of adults were taken by using Nikon D90 digital camera with Nikon MICRO NIKKOR 105 mm lens, and the genitalic figures were made by hand drawing under Motic SMZ168 stereo microscope. The terminology of the genitalia generally follows Aspöck and Aspöck (2008).

Taxonomy

Genus Dilar Rambur

Dilar Rambur, 1838: 9. Type species: Dilar nevadensis Rambur, 1838 (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. Didar Navás, 1913: 6. An incorrect subsequent spelling of Dilar. Lider Kuwayama, 1962: 376. An incorrect subsequent spelling of Lidar.

Diagnosis. A preliminary diagnosis of antennae and wings has been given in Zhang et al. (2014)1 and is not repeated here. Male ninth tergite in dorsal view with a truncate or arcuate anterior incision and a deeply V- or U- shaped posterior incision, leaving a pair of broad hemitergites, which are obtuse distally and densely haired. Male ninth sternite generally much shorter than ninth tergite. Male ectoproct highly specialized, largely covered by ninth tergite, without callus cerci and any large setae, posteroventrally with a pair of bifid unguiform projections and a pair of short, feebly sclerotized, digitiform projections. Male gonocoxite complexes 9, 10 and 11 comprising two pairs of sclerites (i.e. ninth and tenth gonocoxites) and a transverse sclerite (i.e. gonarcus = eleventh gonocixites); gonarcus laterally connecting to bases of ninth gonocoxites. Hypandrium internum generally trapezoidal, with lateral margins slightly arcuate.

Distribution. This genus ranges from northern Africa, through Europe, to Asia, and has been recorded from the following countries: Afghanistan, Algeria, China, France, Greece, India, Iran, Italy, Japan, Korea, Kyrgyzstan, Lebanon, Malaysia, Nepal, Pakistan, Portugal, Russia, Spain, Sri Lanka, Tajikistan, Thailand, Turkey, Turkmenistan, Vietnam (Oswald 1998; Oswald & Schiff 2001; Yang 1999, 2001; Aspöck et al. 2001; Zhang et al. 2014).

Key to males of Dilar from Tibet

1. Forewing with many dark stripes mostly connected with each other (Fig. 3); male tenth gonocoxites inflated medially in dorsal view (Fig. 12)...... D. tibetanus Yang - Forewing with scattered small markings, which are isolated from each other (Figs. 1, 2); male with tenth gonocoxites not inflated medially in dorsal view (Figs. 4, 8) ...... 2 2. Male tenth gonocoxites slenderly elongate in dorsal view; gonarcus slenderly beam-shaped in dorsal view (Fig. 4)...... D. geometroides Aspöck & Aspöck - Male tenth gonocoxites rather short in dorsal view; gonarcus with a pair of elongate spinous projections extended laterally in dorsal view (Fig. 8) ...... D. harmandi (Navás)

1: In connection with our previous paper on Chinese Dilaridae (Zhang et al. 2014) we have to correct a mistakable statement: The redescription and the drawings of the male genitalia of Dilar septentrionalis published by Monserrat (1988) are adequate and absolutely correct, of course. We want to apologize.

Dilar geometroides Aspöck & Aspöck, 1968: 15. Type locality: Nepal (Janakpur). Dilar aspersus Yang, 1988: 197. Type locality: China (Tibet: Linzhi). syn. nov.

Diagnosis. This species is characterized by the forewings with numerous pale brown markings, and the male complexes of gonocoxites 9, 10, and 11 with rather inflated ninth gonocoxites and slenderly elongate tenth gonocoxites, which are bifid at tip. Description (Based on the larger specimens; see remarks). Male. Body length 4.0–7.0 mm; forewing length 10.0–12.0 mm, hindwing length 8.0–10.0 mm. Head brown, with pale yellow setose tubercles. Compound eyes blackish brown. Antenna with ca. 28 segments, yellowish brown, pedicel with brown annular stripes, flagellum unipectinate on most flagellomeres, medial branches much longer than those on both ends, longest branch nearly 3.0 times as long as relevant flagellomere, but branch of 1st flagellomere short and dentate, distal seven flagellomeres simple. Prothorax pale yellowish brown, pronotum brown, with anterior margin and posterolateral corners yellow, medially with a pair of ovoid markings; mesothorax pale yellowish brown, mesonotum dark brown on anterior and lateral margins; metanotum pale yellowish brown, slightly darker on lateral margins. Legs brown, femora blackish brown at tip. Wings hyaline, slightly smoky brown. Forewing ~2.4 times as long as wide, with numerous pale brown spots, arranging irregularly, proximal spots slightly darker, an immaculate area present distal to median nygma; two nygmata present on proximal and median portion of forewing (Fig. 1). Hindwing ~2.2 times as long as wide, much paler than forewing; one nygma present at middle. Veins pale brown. Forewing with trichosors present along wing margin between R and CuA; costal crossveins simple, but occasionally forked; Sc just touching R in pterostigmatic region, terminally leaving several weak veinlets; Rs with four main branches; MA fused with R at wing base, proximally with no crossvein connecting to MP; MP with two main branches; two gradate crossveins present at middle. Hindwing with trichosors present along wing margin between R and CuA; Rs with four main branches. Abdomen yellowish brown, pregenital segments dorsally dark brown. Ninth tergite in dorsal view with an arcuate anterior incision, a nearly U-shaped posterior incision and an elongate dorsoprocessus, leaving a pair of broad hemitergites, which are obtuse distally and densely haired (Fig. 4); in lateral view broad, with straight ventral margin and arcuate posterior margin (Fig. 6). Ninth sternite well-developed, slightly shorter than ninth tergite, arcuately convex posteriad in ventral view (Fig. 5). Ectoproct in dorsal view with a shallowly arcuate anterior incision, posterodorsally with a pair of ventrally curved unguiform projections and a shallowly arcuate anterior incision, posteroventrally with a pair of nearly ovoid flattened projections, a pair of bifid unguiform projections and a feebly sclerotized, digitiform projection (Fig. 4). Ninth gonocoxites inflated, nearly oblong, in dorsal view with unguiform tip; tenth gonocoxites slenderly elongate, incurved, with bifid unguiform tip, submedially with a lobe connecting to ninth gonocoxite; eleventh gonocoxites (= gonarcus) slenderly beam-shaped in dorsal view (Fig. 4), laterally connecting to bases of ninth gonocoxites. Hypandrium internum nearly trapezoidal, with lateral margins slightly arcuate (Fig. 5). Female. Unknown. Materials examined. 12♂ (Paratypes of D. aspersus), CHINA: Tibet, Linzhi [29°39′N, 94°21′E], 3050 m, 8.VI.1978, Fasheng Li (CAU); 1♂, CHINA: Tibet, Naidong [29°13′N, 91°45′E], 3553 m, 19.VI.1981, Tailu Chen (CAU); 1♂, NEPAL: Terhathum, Phalambung [a locality of Kosi State, 27°11′N, 87°30′E], 1760 m, 25.IV.1962, T. Yasuda (NSMT). Material previously studied (i.e. type material of original description (Aspöck & Aspöck 1968): 28♂, NEPAL, Sete [ca. 27°35′N 86°28′E], 2500–3500 m and Tampa Khosi valley [ca. 27°38′N 86°27′E], 2600 m ( majority of the specimens in ZSM, partly in HUAC). Distribution. China (Tibet) (new country record), Nepal (Janakpur, Kosi). Remarks. This species was previously recorded only from Nepal (Aspöck & Aspöck 1968). It appears to be closely related to Dilar tibetanus Yang by the similar male gonocoxite complexes 9, 10 and 11 with inflated, nearly oblong ninth gonocoxites, but it can be easily separated from D. tibetanus by the much paler forewings and the male gonocoxites complexes 9, 10, and 11 with slenderly elongate tenth gonocoxites, which is bifid at tip. In D. tibetanus, the tenth gonocoxites are spindled anteriorly and posteriorly, with median portion inflated.

DILAR FROM TIBET Zootaxa 3878 (6) © 2014 Magnolia Press · 553 FIGURES 1–3. Adults of Dilar spp. 1. D. geometroides Aspöck & Aspöck, male from Tibet; 2. D. harmandi (Navás), male from Tibet; 3. D. tibetanus Yang, male holotype. Scale bars: 1.0 mm.

554 · Zootaxa 3878 (6) © 2014 Magnolia Press ZHANG ET AL. FIGURES 4–7. Dilar geometroides Aspöck & Aspöck. 4. Male genitalia, dorsal view; 5. Male genitalia, ventral view; 6. Male genitalia, lateral view; 7. Male ectoproct, caudal view. e: ectoproct; g: gonarcus = eleventh gonocoxites; gx9: ninth gonocoxite; gx10: tenth gonocoxite; hi: hypandrium internum; S9: sternite 9; T9: tergite 9. Scale bars: 0.5 mm.

D. geometroides shows a remarkable variability, which is characterized by large-sized dark specimens with forewing length 10–12 mm on one hand and small-sized pale specimens with forewing length 7–8 mm on the other hand (Figs. 16 and 17). The Chinese specimens apparently have much narrower forewings than those from Nepal. The variability of the genital sclerites of the males is hardly commensurable quantitatively due to their complexity. The presently available material does not allow to differentiate whether it reflects a clinal phenomenon, polymorphism or character displacement. Dilar aspersus Yang was originally described in a revision of Neuroptera from Tibet by Yang (1988) based on only external morphology, e.g. the marking pattern of mesothorax and the dense small spots on forewings. However, after examining the holotype of D. aspersus and several specimens of D. geometroides from Nepal, we

DILAR FROM TIBET Zootaxa 3878 (6) © 2014 Magnolia Press · 555 found that the male genitalia of D. aspersus is the same as D. geometroides. Therefore, we treat D. aspersus as a junior synonym of D. geometroides.

Dilar harmandi (Navás) (Figs. 2, 8–11, 18, 19)

Nepal harmandi Navás, 1909: 661. Type locality: India (West Bengal).

Diagnosis. This species is characterized by the forewings with several scattered yellowish brown spots, the male complexes of gonocoxites 9, 10, and 11 with rather short tenth gonocoxites, and the gonarcus (= gonocoxites 11) with a pair of elongate spinous projections extended laterally connecting to middle part of ninth gonocoxites. Description (Based on the larger specimens; see remarks). Male. Body length 7.0 mm; forewing length 9.0–10.0 mm, hindwing length 8.5 mm. Head brown, with pale yellowish brown setose tubercles. Compound eyes blackish brown. Antenna with ca. 27 segments, dark brown, pedicel with brown annular stripes, flagellum unipectinate on most flagellomeres, medial branches much longer than those on both ends, longest branch nearly 3.0 times as long as relevant flagellomere, but branch of 1st flagellomere short and dentate, distal seven flagellomeres simple. Prothorax brown, pronotum dark brown, with anterior margin and posterolateral corners slightly paler, medially with a pair of ovoid markings; mesothorax yellowish brown, mesonotum blackish brown, medially with a pair of pale yellow markings; metanotum blackish brown, pale yellow on anteromedian area. Legs brown, femora blackish brown at tip. Wings dirty yellow, with several scattered brown spots. Forewing ~2.2 times as long as wide, with several small brown spots, most of which are concentrated between Sc and Rs, arranging irregularly, a big brown spot present around median nygma; three nygmata present on proximal and median portion of forewing (Fig. 2). Hindwing ~2.1 times as long as wide, slightly smoky brown; one nygma present at middle. Veins brown, crossveins much paler than longitudinal veins. Forewing with trichosors present along wing margin between R and CuA; costal crossveins simple, but occasionally forked; Sc just touching R in pterostigmatic region, terminally leaving several weak veinlets; Rs with three main branches; MA fused with R at wing base, proximally with no crossvein connecting to MP; MP with two main branches; two gradate crossveins present at middle. Hindwing with trichosors present along wing margin between R and CuA; Rs with four main branches. Abdomen yellowish brown, pregenital segments dorsally dark brown. Ninth tergite in dorsal view with an arcuate anterior incision, a nearly U-shaped posterior incision and an elongate dorsoprocessus, leaving a pair of broad hemitergites, which are obtuse distally and densely haired (Fig. 8); in lateral view broad, with straight ventral margin and arcuate posterior margin (Fig. 10). Ninth sternite much shorter than ninth tergite, arcuately convex posteriad in ventral view (Fig. 9). Ectoproct in dorsal view with an arcuate anterior incision, posterodorsally with a pair of ventrally curved unguiform projections, posteroventrally with a pair of nearly ovoid flattened projections, a pair of bifid unguiform projections and a feebly sclerotized, digitiform projections (Fig. 8). Ninth gonocoxites slenderly elongate, with anterior half broadly spoon-shaped and with posterior half slightly incurved and nearly unguiform at tip in dorsal view; tenth gonocoxites rather short, nearly rod-shaped; eleventh gonocoxites (= gonarcus) beam-shaped in dorsal view (Fig. 8), with a pair of elongate spinous projections extended laterally connecting to middle part of ninth gonocoxites. Hypandrium internum nearly trapezoidal, with lateral margins slightly arcuate (Fig. 9). Female. Unknown. Materials examined. Holotype ♂, INDIA: West Bengal, Darjeeling [27°02′N, 88°20′E], 1890, Harmand leg. (MNHN). 1♂, CHINA: Tibet, Yadong, Mt. Naituilashan [27°24′N, 88°51′E], 3650 m, 18.VII.2013, Meicai Wei & Gengyun Niu (CAU); 1♂, CHINA: Tibet, Zhangmu, Lixin [27°59′N, 85°58′E], 2500 m, 24.VI.1975, Xuezhong Zhang (CAU); 1♂, NEPAL: Taplejung Walungchung Gola [a locality of Mechi State, 27°37′N, 87°46′E], 3350 m, 18.VI.1962, T. Yasuda (NSMT); 1♂, NEPAL: Taplejung Walungchung Gola, 3310 m, 11.VII.1962, T. Yasuda (NSMT); 2♂, NEPAL: Prov. Nr.3 East Bujan, Dudh Kosi Tal, 2900 m, 18.–19.VII.1964, R. Remane; 7♂, NEPAL: Prov. Nr.3 East Dudh Kosi Tal unter Thangpoche, 3400 m, 4.VII.1964, R. Remane; 16♂, NEPAL: Prov. Nr.3 East Sete, 3000–3600 m, 23.VII.1964, R. Remane; 11♂, NEPAL: Prov. Nr.3 East Surkaya, 2600–3000 m, 18.VII.1964, R. Remane; 2♂, NEPAL: Prov. Nr.3 East Lamjura Pass, 3200–3600 m, 1.VIII.1964, R. Remane; 2♂, NEPAL: Prov.

556 · Zootaxa 3878 (6) © 2014 Magnolia Press ZHANG ET AL. Nr.3 East Jambesi, 2900–3300 m, 23.VI.1964, R. Remane; 5♂, NEPAL: Prov. Nr.1 East Phulchauki, 2500–2800 m, 30.V.1964, R. Remane; 1♂, NEPAL: Khumbu, Khumdzung, 3900 m, 11.VII.1962, Ebert-Falkner (majority of the Nepalese specimens collected by R. Remane in ZSM, partly in HUAC).

FIGURES 8–11. Dilar harmandi (Navás). 8. Male genitalia, dorsal view; 9. Male genitalia, ventral view; 10. Male genitalia, lateral view; 11. Male ectoproct, caudal view. Scale bars: 0.5 mm.

Distribution. China (Tibet) (new country record), India (West Bengal), Nepal (Mechi, Bagmati, Gandaki) (new country record). Remarks. Most specimens of D. harmandi can be easily distinguished from the other two Dilar species from Tibet by the dirty yellow wings with several scattered brown spots. However, the variability in the general appearance of D. harmandi is as bewildering as that of D. geometroides discussed above. D. harmandi comprises large dark specimens (forewing length 9–10 mm) and smaller and paler specimens (forewing length ca. 7 mm) (Figs. 18 and 19). The genital sclerites of the males don’t show significant intraspecific differences, however, their

DILAR FROM TIBET Zootaxa 3878 (6) © 2014 Magnolia Press · 557 complexity prevents them from being efficiently comparable. Based on the male genitalia, D. harmandi differs from the other dilarid species found in Tibet by the male complexes of the gonocoxites 9, 10, and 11, with rather short tenth gonocoxites and the eleventh gonocoxites (= gonarcus) with a pair of elongate spinous projections extended laterally connecting to the middle part of ninth gonocoxites.

Dilar tibetanus Yang (Figs. 3, 12–15)

Dilar tibetanus Yang, 1987: 197. Type locality: China (Xizang). Dilar pumilus Yang, 1988: 197. Type locality: China (Xizang). Dilar pusillus Yang, 1992: 379. Type locality: China (Xizang). syn. nov.

Diagnosis. This species is characterized by the forewings with many dark stripes mostly connected with each other, and by the male complexes of the gonocoxites 9, 10, and 11 with suboblong ninth gonocoxites and with medially inflated tenth gonocoxites which is spindled anteriorly and posteriorly. Description. Male. Body length 2.8–4.5 mm; forewing length 5.0–8.0 mm, hindwing length 4.2–7.0 mm. Head dark brown, with pale yellow setose tubercles. Compound eyes blackish brown. Antenna with ca. 25 segments, yellowish brown, pedicel with brown annular stripes, flagellum unipectinate on most flagellomeres, medial branches much longer than those on both ends, longest branch nearly 4.0 times as long as relevant flagellomere, but branch of 1st flagellomere short and dentate, distal eight flagellomeres simple. Prothorax pale yellowish brown, pronotum brown, with anterior margin and posterolateral corners yellow, medially with a pair of ovoid markings; mesothorax pale yellowish brown, mesonotum dark brown on anterior and lateral margins; metanotum pale yellowish brown, slight darker on lateral margins. Legs yellowish brown, femora blackish brown at tip. Wings yellowish brown. Forewing ~2.1 times as long as wide, with numerous dark brown stripes, proximal stripes slightly darker, most markings on distal half connected with each other, generally arranging as transversely arcuate stripes, a broad brown spot present around median nygma; three nygmata present on proximal and median portion of forewing (Fig. 3). Hindwing ~2.0 times as long as wide, pale brown; one nygma present at middle. Veins pale brown. Forewing with trichosors present along wing margin between R and CuP; costal crossveins simple, but occasionally forked; Sc just touching R in pterostigmatic region, terminally leaving several weak veinlets; Rs with three main branches; MA fused with R at wing base, proximally with no crossvein connecting to MP; MP with two main branches; two gradate crossveins present at middle, inconspicuous due to pale coloration. Hindwing with trichosors present along wing margin between R and CuA; Rs with four main branches. Abdomen yellowish brown, pregenital segments dorsally dark brown. Ninth tergite in dorsal view with an arcuate anterior incision, a nearly U-shaped posterior incision and an elongate dorsoprocessus, leaving a pair of broad hemitergites, which are obtuse distally and densely haired (Fig. 12); in lateral view broad, with straight ventral margin and arcuate posterior margin (Fig. 14). Ninth sternite much shorter than ninth tergite, arcuately convex posteriad in ventral view (Fig. 13). Ectoproct in dorsal view with a deeply arcuate anterior incision, posterodorsally with a pair of ventrally curved unguiform projections, posteroventrally with a pair of nearly ovoid flattened projections, a pair of bifid unguiform projections and a feebly sclerotized, digitiform projection (Fig. 12). Ninth gonocoxites inflated, suboblong in dorsal view with blunt tip; tenth gonocoxites spindled anteriorly and posteriorly, with median portion inflated (Fig. 12); eleventh gonocoxites (= gonarcus) slenderly beam-shaped in dorsal view, laterally connecting to bases of ninth gonocoxites. Hypandrium internum nearly trapezoidal, with lateral margins slightly arcuate (Fig. 13). Female. Unknown. Materials examined. Holotype ♂, CHINA: Tibet, Yigong [30°16′N, 94°49′E], 2300 m, 15.VI.1978, Fasheng Li (CAU). Paratypes 3♂, same data as holotype (CAU); 1♂, CHINA: Tibet, Yigong [30°16′N, 94°49′E], 2300 m, 26.VIII.1983, Yinheng Han (CAU); 2♂, CHINA: Tibet, Zhamu [29°20′N, 91°49′E], 2700 m, 6.VI.1978, Fasheng Li (CAU); 1♂, CHINA: Tibet, Bomi [29°51′N, 95°46′E], 3050 m, 13.VII.1978, Fasheng Li (CAU). 2♂, CHINA: Tibet, Bomi, Mt. galonglashan [29°51′N, 95°46′E], 3026 m, 13.VII.2013, Xiaoyan Liu (CAU); 1♂, CHINA: Tibet, Linzhi [30°01′N, 95°58′E], 2030 m, 12.VI.2009, Meicai Wei (CAU).

558 · Zootaxa 3878 (6) © 2014 Magnolia Press ZHANG ET AL. FIGURES 12–15. Dilar tibetanus Yang. 12. Male genitalia, dorsal view; 13. Male genitalia, ventral view; 14. Male genitalia, lateral view; 15. Male ectoproct, caudal view. Scale bars: 0.5 mm.

Distribution. China (Tibet). Remarks. This species can be distinguished from the other Dilar species from Tibet by the dark wings with many dark stripes on the forewings and by the male complexes of the gonocoxites 9, 10 and 11 with tenth gonocoxites spindled anteriorly and posteriorly, with median portion inflated. Dilar pusillus Yang was originally described based on a several specimens from Tibet, and considered to be distinguished from D. tibetanus by the small body-size, the different marking pattern on head (Yang 1988). However, after examining the holotype of D. pusillus, we found the male genitalia of D. pusillus are also of the same shape as D. tibetanus. Therefore, we treat D. pusillus as a junior synonym of D. tibetanus.

DILAR FROM TIBET Zootaxa 3878 (6) © 2014 Magnolia Press · 559 FIGURES 16–19. Individual variations of Dilar spp. 16. D. geometroides Aspöck & Aspöck, small-sized male from Nepal; 17. same species, large-sized male from Nepal; 18. D. harmandi (Navás), large-sized male from Nepal with dark wings; 19. same species, small-sized male from Nepal with bright wings. Scale bars: 1.0 mm.

Discussion

The present study of species of the genus Dilar from Tibet and the previous paper on species from northern China (Zhang et al. 2014) are contributions to a planned series of revisions of the genus Dilar in China. Dilar is a highly diverse genus in the family Dilaridae having complex structures of male genitalia. However, the male genital sclerites of the genus Dilar can only be understood in the context of knowledge about the other genera, especially of Nallachius (subfamily Nallachiinae). The complex structure which had all along been called “supraanale” (Aspöck et al. 1980, Monserrat 1988, Oswald & Schiff 2001) clearly represents the ectoptoct (= tergites 10+11). This becomes evident when comparing Dilar with Nallachius (Minter 1986; Penny 1994; Aspöck & Aspöck 2008; Machado & Rafael 2010). The striking variability of size, shape, and coloration known from several dilarid species is truly bewildering. This might be a general phenomenon in the Dilaridae. The biology of the Dilaridae is largely unknown, however, unusually high numbers of larval stages have been observed under laboratory conditions in Nallachius (MacLeod & Spiegler 1961). Tibet is considered to be one of the most active geological regions and the most sensitive and richest regions in biological diversity (Yang et al. 2004). The three Dilar species herein redescribed show diverse forewing marking patterns, varying from scattered dotted to densely banding patterns. However, these Tibetan species have considerable interspecific similarity in the male genitalia, e.g. ninth tergite with similar dorsoprocessus and ectoproct posterodorsally with a pair of ventrally curved unguiform projections – a phenomenon which is apparently due to a young radiation. Of the three Tibetan Dilar species, two species are also distributed in Nepal, indicating close relationships between the fauna of Dilaridae from southeastern Tibet and Nepal. The uplifting of Himalayas was one of the significant factors promoting the speciation and shaping the diverse fauna of East Asia.

560 · Zootaxa 3878 (6) © 2014 Magnolia Press ZHANG ET AL. Up to now, seven species of Dilaridae have been described from the areas around the Himalayas (including Nepal, northeastern India, and Tibet), and all the complexes of male gonocoxite 9, 10 and 11 of these species are obviously different from each other (Aspöck & Aspöck 1968; McLachlan 1869; Monserrat 1989; Nakahara 1963; Navás 1909). Some of the species need revision, however, more species of this enigmatic lacewing family remain to be discovered in the future from this area.

FIGURE 20. Geographic distribution of the species of Dilar from Tibet, China. ▲: Dilar geometroides Aspöck & Aspöck; ●: Dilar harmandi (Navás); ■: Dilar tibetanus Yang.

Acknowledgements

We thank Dr. Xiaoyan Liu (Wuhan) for collecting specimens from Tibet. We also thank Dr. Utsugi Jinbo (NSMT, Tsukuba), Dr. Masaaki Tomokuni (NSMT, Tsukuba), Dr. Meicai Wei (Changsha), and Dr. Gengyun Niu (Changsha) for processing the loan of specimens in their collections. We gratefully thank Dr. Heinz Wundt (ZSM, Munich) for the loan of the Dilaridae of the ZSM, and Dr. Jean Legrand (MNHN, Paris) for the loan of the holotype of D. harmandi. This research was supported by the National Natural Science Foundation of China (Nos. 31322051 and 31320103902), the National Key Basic Research Program of China (973 Program) (No. 2013CB127601), and the Foundation for the Author of National Excellent Doctoral Dissertation of PR China (No. 201178).

References

Aspöck, H. & Aspöck, U. (1968) Zwei weitere neue Spezies des Genus Dilar Rambur (Neuroptera, Planipennia) aus Asien. (Vorläufige Mitteilung). Entomologisches Nachrichtenblat, Wien, 15, 3–6.

DILAR FROM TIBET Zootaxa 3878 (6) © 2014 Magnolia Press · 561 Aspöck, H., Aspöck, U. & Hölzel, H. (unter Mitarbeit von H. Rausch) (1980) Die Neuropteren Europas. Goecke und Evers, Krefeld, 495 pp. [355 pp] Aspöck, H., Hölzel, H. & Aspöck, U. (2001) Kommentierter Katalog der Neuropterida (Insecta: Raphidioptera, Megaloptera, Neuroptera) der Westpaläarktis. Denisia, 2, 1–606. Aspöck, U. & Aspöck, H. (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology, 33, 97–127. http://dx.doi.org/10.1111/j.1365-3113.2007.00396.x Huang, F.S. & Han, Y.H. (1988) Insects in Mt. Namjagbarwa Region of Xizang. In: Huang, F.S. (Ed.), Insects of Mt. Namjagbarwa Region of Xizang. Science press, Beijing, pp. 1–17. [in Chinese] Machado, R.J.P. & Rafael, J.A. (2010) Two new species of Dilaridae (Insecta: Neuroptera) with additional notes on Brazilian species. Zootaxa, 2421, 61–68. McLachlan, R. (1869) On a neuropterous insect from N. W. India, belonging to the genus Dilar. Entomologist's Monthly Magazine, 5, 239–240. MacLeod, E.G. & Spiegler, P.E. (1961) Notes on the larval habitat and developmental peculiarities of Nallachius americanus (McLachlan) (Neuroptera: Dilaridae). Proceedings of the Entomological Society of Washington, 63, 281–286. Minter, L.R. (1986) The first record of Dilaridae (Neuroptera) from the Afrotropical region. Journal of the Entomological Society of Southern Africa, 49 (1), 87–94. Monserrat, V.J. (1988) Revisión de la obra de L. Navas, I: EL genero Dilar Rambur, 1842 (Neuropteroidea, Planipennia: Dilaridae). Neuroptera International, 5 (1), 13–23. Monserrat, V.J. (1989) Algunos Neurópteros del Museo de Basilea (Insecta, Neuropteroidea, Planipennia). Entomologica Basiliensia, 13, 417–428. Nakahara, W. (1963) A remarkable new dilarid from India (Neuroptera). Kontyû, 31, 77–78. Navás, L. (1909) Monografía de la familia de los Diláridos (Ins. Neur.). Memorias de la Real Academia de Ciencias y Artes de Barcelona, 3 (7), 619–671. Oswald, J.D. (1998) Annotated catalogue of the Dilaridae (Insecta: Neuroptera) of the world. Tijdschrift voor Entomologie, 141, 115–128. http://dx.doi.org/10.1163/22119434-99900008 Oswald, J.D. & Schiff, N.M. (2001) A new species of the genus Dilar Rambur (Neuroptera: Dilaridae) from Borneo. Proceedings of the Entomological Society of Washington, 103, 74–80. Oswald, J.D. (2013) Dilaridae. Neuropterida Species of the World. Version 2.0. Available from: http://lacewing.tamu.edu/ Species-Catalogue/ (Accessed 14 September 2013) Penny, N.D. (1994) A new species of Nallachius (Neuroptera: Dilaridae) from Costa Rica. Pan-Pacific Entomologist, 70 (4), 309–312. Yang, C.K. (1987) Neuroptera. In: Zhang, S. (Ed.), Xizang nong ye bing chong ji za cao. Vol.1. Xizang Renmin Press House, Xizang, pp. 191–220. Yang, C.K. (1988) Neuroptera: Osmylidae, Dilaridae, Hemerobiidae, Chrysopidae, Mantispidae, Myrmeleontidae, Ascalaphidae, Corydalidae. In: Huang, F.S., Wang, P.Y., Yin, W.Y., Yu, P.Y., Lee, T.S., Yang, C.K. & Wang, X.J. (Eds.), Insects of Mt. Namjagbarwa region of Xizang. Science press, Beijing, pp. 193–216. Yang, C.K. (1992) Neuroptera. In: Chen, S. (Ed.), Insects of the Hengduan Mountains Region. Science press, Beijing, pp. 438–454. Yang, C.K. (1999) Dilaridae. In: Huang, B.K. (Ed.), Fauna of Insects Fujian Province of China. Fujian Science and Technology Press, Fuzhou, pp. 94–95. Yang, C.K. (2001) Neuroptera: Mantispidae and Dilaridae. In: Wu, H. & Pan, C.W. (Eds.), Insects of Tianmushan National Nature Reserve. Science press, Beijing, pp. 305–307. Yang, X.K., Huang, F.S. & Yao, J. (2004) Marvelous insect kingdom—the Great Yarlung Zangbo Canyon. In: Yang, X.K. (Ed.), Insect of the Great Yarlung Zangbo Canyon of Xizang, China. China Science and Technology Press, Beijing, pp. 1–17. [in Chinese] Zhang, W., Liu, X.Y., Aspöck, U. & Aspöck, H. (2014) Revision of Chinese Dilaridae (Insecta: Neuroptera) (Part I): Species of the genus Dilar Rambur from northern China. Zootaxa, 3753 (1), 10–24. http://dx.doi.org/10.11646/zootaxa.3753.1.2