Dtsch. Entomol. Z. 56 (2) 2009, 317–321 / DOI 10.1002/mmnd.200900029

Inocellia elegans sp. n. (Raphidioptera, Inocelliidae) – A new and spectacular from China

Xingyue Liu*,1,2, Horst Aspck3, Ding Yang1 and Ulrike Aspck4

1 Department of Entomology, China Agricultural University, Beijing 100193, China 2 Department of Biology, Tokyo Metropolitan University, Minamiohsawa 1-1, Hachioji, Tokyo 192-0397, Japan 3 Department of Medical Parasitology, Medical University of Vienna, Kinderspitalgasse 15, A-1095 Vienna, Austria 4 Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, A-1010 Vienna, Austria

Abstract

Received 3 June 2009 A new snakefly species of Inocelliidae, Inocellia elegans sp. n., from Guizhou Pro- Accepted 10 June 2009 vince, China, is described and illustrated. The appearance of this species is rather spec- Published 11 November 2009 tacular and differs from all other known species of Raphidioptera by the pigmented wings. The systematic position of this species is discussed. Key Words

New species Zoogeography Oriental Region

Introduction et al. 1991). After the revision of this by H. As- pck et al. (1991), Yang (1999) described two new spe- The snakefly genus Inocellia Schneider, 1843 is the lar- cies, Inocellia fujiana Yang, 1999 and Inocellia as- gest genus of the family Inocelliidae. It is widespread pouckorum Yang, 1999, from China, whereby the latter in the Palaearctic region and the transgressional areas species was named only on the basis of the re-illustra- between eastern Palaearctic and northern part of Orien- tion and re-description of I. sinensis made by U. Aspck tal regions. The genus is characterized by the stylus on & H. Aspck (1980) so that its validity is doubtful. the subdistal portion of the male ninth gonocoxite, the Therefore, although 10 extant species of Inocellia are bristle tufts on the inner side of the male ninth gono- known, the actual number of valid species might be coxite and the endophallus, and the feebly sclerotized around seven. foliate pseudostyli (basal parts of the gonapophyses 9). Considering the geographical distribution, except for In the monograph of H. Aspck et al. (1991), six I. crassicornis which is widespread in the Palaearctic species were listed under the genus Inocellia and sepa- realm, the other Inocellia species are restricted to com- rated into two species groups, namely the Inocellia paratively small areas and disjunctly distributed from crassicornis group (Inocellia crassicornis (Schummel, each other in Afghanistan, Pakistan, northern India, 1832), Inocellia taiwana H. Aspck & U. Aspck, Bhutan, Chinese mainland, Taiwan, and Japan, showing 1985, and Inocellia japonica Okamoto, 1917) and the a tendency to diversify in Asia. On the mainland China, Inocellia fulvostigmata group (Inocellia fulvostigmata there are only three species currently known from quite U. Aspck & H. Aspck, 1968, Inocellia bhutana a few localities in the eastern part. However, it should H. Aspck, U. Aspck & Rausch, 1991, and Inocellia be noticed that, judging from the rather limited amount sinensis Navs, 1936). Additionally, two species, Ino- of available materials of most species, specimens of In- cellia frigida Navs, 1915 and Inocellia rossica Navs, ocellia are rarely encountered in the field probably due 1916, were considered as nomina dubia, and they are to the low population density, so that more unknown most probably synonyms of I. crassicornis (H. Aspck species are likely to be found in China and its adjacent

* Corresponding author, e-mail: [email protected]

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 318 Xingyue Liu et al.: New snakefly from China (Raphidioptera, Inocelliidae) areas if a long-term intensive investigation is to be car- hindwing cryptic. Male ninth gonocoxite shell-like, ried out. much wider than long in the I. crassicornis group, In the present study, a spectacular Inocellia species, while much longer than wide in the I. fulvostigmata collected in the national nature reserve of Mt. Fanjing- group; inner side with a stylus near subdistal portion, shan, Guizhou Province, is described and illustrated as and usually bearing some bristle tufts. Pseudostyli (ba- new to science. The systematic position of the new spe- sal parts of gonapophyses 9) feebly sclerotized, gener- cies is discussed. ally narrow and foliate, nearly connected with each other at middle. Parameres (gonocoxite-gonapophyses- gonostyli complex 10) unpaired, proximal portion flat- Materials and methods tened, with a slender distal projection. Gonarcus (gono- The specimen upon which the present study is based was occasionally coxites 11) generally shield-like, sometimes with pro- collected through sweeping net by beating trees. After collection, it jections from its posterior surface. Endophallus short, was directly preserved in 70% alcohol and is currently deposited in mostly with some bristles or bristle tufts. Female se- the Entomological Museum of China Agricultural University (CAU), venth sternum broad, posteriorly produced more or less, Beijing. Genitalic preparations were made by clearing the apex of the with posterior margin truncate or concaved; subgenital abdomen in a cold, saturated KOH solution for 6–8 h. After rinsing plate reduced into a small ovoid sclerite or much more the KOH with acetic acid and water, the apex of the abdomen was modified into an irregularly shaped plate. transferred to glycerin for further dissection and examination. The ter- minology of the genitalia generally follows that of H. Aspck et al. Distribution. Europe (Austria, Czech Republic, Eur- (1991) and U. Aspck & H. Aspck (2008). opean part of Russia, Finland, Germany, Netherlands, Poland, Sweden); Asia (Afghanistan, Armenia, Bhutan, Taxonomy China, India, Japan, Mongolia, North Korea, Pakistan, Russia Far East) (H. Aspck et al. 1991, 2001). Genus Inocellia Schneider Remarks. The close affinity between Inocellia and Inocellia Schneider, 1843: 84. Type species: Raphidia crassicornis Parainocellia was proposed due to the presence of the Schummel, 1832 (designated by monotypy). bristles or bristle tufts on the inner side of the male Diagnosis. Adult generally blackish brown with yellow- ninth gonocoxite, as well as on the endophallus (H. As- ish thoracic and abdominal markings, and sometimes pck et al. 1991). Nonetheless, Inocellia can be easily with pale markings on head. Antennae and legs usually separated from Parainocellia by the feebly sclerotized pale yellow or yellowish brown. Media anterior in the and foliate pseudostyli of the latter.

Figure 1. Habitus of Inocellia elegans sp. n. (holotype male). Scale bars ¼ 1.0 mm.

museum-dez.wiley-vch.de # 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Dtsch. Entomol. Z. 56 (2) 2009, 317–321 319

Inocellia elegans sp. n. ish transverse marking on posterior margin of scutel- Figures 1–8 lum. Legs yellow with brownish setae except coxae brown; apices of tibiae slightly darker. Wings hyaline Diagnosis. Wings hyaline with distal 1/3 brown; male but distal 1/3 tinged with brown, pterostigma and veins ninth gonocoxite elongated, nearly twice as long as brown. Anterior branch of Rs with one trifurcated vein ninth tergum; endophallus with two pairs of bristle tufts and one simple vein running to wing margin. and eight strong bristles. Abdomen gradually enlarged posteriorly from 5th ab- Male. Body length 14.0 mm; forewing length dominal segment, with each of 5th to 8th segments sub- 13.0 mm, hindwing length 11.5 mm. trapezoidally widened posteriorly; coloration generally Head subquadrate, black, with clypeus pale brown; blackish brown with venter slightly paler; each of 1st to occiput medially with a pair of yellowish brown stripes. 6th pregenital segments with a yellowish transverse Compound eyes blackish brown. Antennal sclerite stripe posteriorly except for 1st tergum entirely blackish brown, scape, and pedicel pale yellow, flagellum brown brown; genital segments blackish brown, ninth tergum with proximal 7–8 segments pale yellow. Mouthparts laterally with posterior margin brownish yellow, ninth reddish brown, mandibles dark brown. gonocoxite entirely brownish yellow, ectoproct with Thorax blackish brown; anterior margin of pronotum posterior margin pale yellow. Ninth tergum (Figs 2–3) yellow; mesonotum and metanotum each with a yellow- in dorsal view nearly rounded, anterior margin slightly

Figures 2–8. Male genitalia of Inocellia elegans sp. n.; 2. Genital segments, lateral view; 3. Genital segments, dorsal view; 4. Gen- ital segments, ventral view; 5. Ninth gonocoxite and internal structures, lateral view; 6. Gonarcus (¼ gonocoxites 11), caudal view; 7. Gonarcus, dorsal view; 8. Parameres (¼ gocoxites-gonapophyses-gonostyli complex 10), dorsal view. Scale bars ¼ 0.5 mm. Ab- breviations: e – ectoproct; ep – endophallus; g – gonarcus; gx – ninth gonocoxite; p – parameres; ps – pseudostyli (¼ gonapophyses 9); s – stylus; S8–9 – eighth and ninth sternum; T8–9 – eighth and ninth tergum.

# 2009 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-dez.wiley-vch.de 320 Xingyue Liu et al.: New snakefly from China (Raphidioptera, Inocelliidae) incised medially, posterior margins medially with sub- cellia and Navs but rather a synplesiomorphy. trapezoidal incision. Ninth sternum (Figs 2, 4) in ventral And although gigantism of Sininocellia gigantos and view broad, anteriorly prominent medially, posterior Inocellia elegans is a derived character, and certainly a margin slightly concaved medially. Ninth gonocoxite rare phenomenon, it is not necessarily a synapomorphy (Figs 2–5) shell-like but strongly elongated, nearly of these taxa but an independent acquisition. The lack twice as long as ninth tergum, slightly narrowed toward of the Media anterior in the hindwing of Inocellia is apex with rounded tip; inner side medially bearing a apomorphic. Considering the elongated male ninth go- small subquadrate stylus and a single bristle anterior to nocoxite, I. elegans appears to be related to the species stylus near ventral margin, dorsally with a bristle tuft of the I. fulvostigmata group. Furthermore, the presence directed ventrad near proximal portion and with a row of the hypandrium internum in I. elegans is doubted be- of bristles medially. Parameres (gonocoxite-gonapo- cause it was not found despite careful dissection and physes-gonostyli complex 10) (Figs 5, 8) unpaired, prox- search for it. imal portion flattened, slightly widened ventrad, with Mt. Fanjingshan, the type locality of the new species, ventral margin slightly incised medially; distal projec- is the main mountain of the Wuling Mountain Region tion short, hook-like, and curved dorsad, with a pair of in central China, ranging from 1000 m to 2570 m in al- claws connected to dorsal margin of proximal portion titude, covered by evergreen forest below 1300 m and by a membrane. Gonarcus (gonocoxites 11) (Figs 5–7) mingled with alpine coniferous forest and meadow in shield-like, in lateral view subtrapezoidal, with obtuse the higher areas (Li & Jin 2006). The area of the type distal corners. Endophallus (Figs 4–5) short and rather locality of I. elegans with its high altitudes can be inter- broad, dorsal portion proximally with a pair of bristle preted as a postglacial compensation of ecological re- tufts anteriorly directed dorsad, distal portion laterally quirements of the species (see discussion in Liu et al. sclerotized, bearing a pair of bristle tuft directed poster- (in press)). Mt. Fanjingshan is considered to be a hot- iorly, ventral portion medially with eight strong bristles. spot of biodiversity in China as postglacial refugial area Pseudostyli (gonapophyses 9) (Fig. 4) feebly sclerotized, harboring many endemic organisms (Li & Jin 2006). narrowly foliate, anteriorly curved and almost connected Therefore, I. elegans probably represents an indepen- with each other in ventral view. Ectoproct (Figs 2–3) dent and highly derived lineage within the genus Ino- subquadrate, in lateral view with dorsal margin slightly cellia due to a geographical isolation. elevated. Hypandrium internum lost. Materials examined. Holotype <, CHINA: Guizhou Province, Jiangkou Acknowledgements County, Mt. Fanjingshan, Huguosi, 27550 N, 108410 E, 1158 m, 29.V.2002, Ding Yang (CAU). We are grateful to Prof. Zizhong Li (Guiyang) for his kind invitation Distribution. China (Guizhou). to visit the national nature reserve of the Mt. Fanjing in Guizhou Pro- vince in 2002 and for making arrangements to enable our field trip Etymology. The specific epithet ‘elegans’ refers to the during which the new species described here was discovered. We also spectacular and beautiful appearance of the new species thank Drs. Fumio Hayashi and Yukimasa Kobayashi (Tokyo) for their by the conspicuously pigmented wings. hospitality during the study of the first author in Tokyo Metropolitan University, Japan. Finally, sincere thanks are given to Dr. John Plant (Vienna) for linguistic improvement. This research was supported by Discussion the grant-in-aid for JSPS fellows relating to JSPS Postdoctoral Fellow- ship for Foreign Researchers (No. 20-08417) and by the Initiative Foundation for Research of China Agricultural University Inocellia elegans is the only known snakefly species (No. 10099821). with pigmented wings and can therefore be easily dis- tinguished from all other raphidiopterans – provided that there is no sexual dimorphism and that the hitherto References unknown female is of a similar appearance, in particu- lar with pigmented wings. The placement of this spe- Aspck, H. & Aspck, U. 1985. Inocellia taiwana n. sp. – eine neue cies within Inocellia refers to the presence of the bristle Inocelliiden-Spezies aus Taiwan (Neuropteroidea: Raphidioptera: tufts on several male genitalic structures and the foliate Inocelliidae). Entomologische Zeitschrift mit Insektenbrse 95: 45 48. pseudostyli. However, this species is unique amongst – Aspck, H., Hlzel, H., & Aspck, U. 2001. Kommentierter Katalog the Inocellia species by the large body size and the dis- der Neuropterida (Insecta: Raphidioptera, Megaloptera, Neurop- tinctly broadened abdomen. In this connection it is tera) der Westpalarktis. – Denisia 02: 606 pp; 6 Abb. worth mentioning that another even more gigantic Ino- Aspck, H., Aspck, U. & Rausch, H. 1991. Die Raphidiopteren der celliid, with 21 mm length of forewings, was found in Erde. Eine monographische Darstellung der Systematik, Taxo- Eastern China (Yang 1985): Sininocellia gigantos Yang, nomie, Biologie, kologie und Chorologie der rezenten Raphi- 1985. The only existing specimen, the female holotype, diopteren der Erde, mit einer zusammenfassenden bersicht der fossilen Raphidiopteren (Insecta: Neuropteroidea). 2 Bnde. Goecke was studied by U. Aspck & H. Aspck (1999) who in- & Evers, Krefeld: 730 pp, 550 pp. cluded a discussion of its systematic position. The Aspck, U. & Aspck, H. 1968. Zwei neue Inocelliiden-Spezies aus longitudinal basal piece of the Media anterior in the Asien (Neuroptera, Raphidiodea). – Entomologische Berichte 28: hindwing is not necessarily a synapomorphy of Sinino- 184–189.

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