Inocellia Rara Sp. Nov. (Raphidioptera: Inocelliidae), a New Snakefly

Zootaxa 3753 (3): 226–232 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.3753.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:D79E4111-3409-4A28-A11D-B79EBB90F85C Inocellia rara sp. nov. (Raphidioptera: Inocelliidae), a new snakefly species from Taiwan, with remarks on systematics and biogeography of the Inocelliidae of the island

XINGYUE LIU1, HORST ASPÖCK2 & ULRIKE ASPÖCK 3,4 1Department of Entomology, China Agricultural University, Beijing 100193, China. E-mail: [email protected] 2Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University (MUW), Kinderspitalgasse 15, 1090 Vienna, Austria. E-mail: [email protected] 3Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, A-1010 Vienna, Austria. E-mail: [email protected] 4Department of Evolutionary Biology, University of Vienna, Althanstraße 14, 1090 Vienna, Austria

Abstract

A new species of the snakefly genus Inocellia Schneider, 1843 from Taiwan is described: Inocellia rara sp. nov. It represents the third species in the family Inocelliidae and the first record of the Inocellia fulvostigmata species group from Tai- wan.

Key words: snakefly, Inocellia, new species, Taiwan, China

Introduction

The Inocelliidae is one of the two families of the endopterygotan insect order Raphidioptera (commonly known as snakeflies), which is one of the smallest insect orders comprising altogether about 240 valid species. Hitherto, there were 39 valid species of Inocelliidae in the world (H. Aspöck et al. 2012; Liu et al. 2012a, 2012b; H. Aspöck & U. Aspöck 2013; Liu et al. 2013). It is remarkable that 17 of the species, which represent ~45% of the world’s Inocelliidae, were discovered and described in the past five years, and all of them come from China and Thailand (U. Aspöck et al. 2011; Liu et al. 2009a, 2009b, 2010b, 2010c, 2012a, 2012b). This indicates the presence of a diverse fauna of this family in East and Southeast Asia. Taiwan is the largest island of China and harbours six endemic species of Raphidioptera (H. Aspöck & U. Aspöck 1985; U. Aspöck et al. 2009; Liu et al. 2010a), among which only two species belong to Inocelliidae, namely Inocellia taiwana H. Aspöck & U. Aspöck, 1985 and Inocellia shinohara U. Aspöck, Liu & H. Aspöck, 2009. These two species belong to the Inocellia crassicornis species group, in which the male gonocoxite 9 is wider than long, and they are distributed in adjacent localities of Nantou County, central Taiwan. In the present paper, we describe a new inocelliid species based on a single specimen collected in northern Taiwan in the early 1940s. This new species is the third endemic species of Inocelliidae from Taiwan and represents the first record of Inocellia fulvostigmata species group in this island.

Material and methods

The type specimen of the presently described new species is deposited in the National Institute for Agro- Environmental Sciences, Tsukuba, Japan (NIAES). Genitalic preparations were made by clearing the apex of the abdomen in a cold saturated KOH solution for 6–8 h. After rinsing the KOH with acetic acid and water, the apex of the abdomen was transferred to glycerin for further dissection and examination. The terminology of the genitalia generally follows that of H. Aspöck et al. (1991) and U. Aspöck and H. Aspöck (2008).

226 Accepted by B. Price: 2 Dec. 2013; published: 7 Jan. 2014 Taxonomy

Genus Inocellia Schneider

Inocellia Schneider, 1843: 84. Type species: Raphidia crassicornis Schummel, 1832 (designated by monotypy).

Diagnosis. Adults generally blackish-brown with yellowish thoracic and abdominal markings, and sometimes with pale markings on head. Antennae and legs usually pale yellow or yellowish brown. Media anterior in the hindwing absent. Male gonocoxite 9 dome-like, much wider than long in the I. crassicornis group, while much longer than wide in the I. fulvostigmata group; inner side with a stylus near subdistal portion, and usually bearing some bristle tufts. Pseudostyli (basal parts of gonapophyses 9) feebly sclerotized, generally narrow and foliate, closely approximated to each other in the middle. Parameres (gonocoxite-gonapophyses-gonostyli complex 10) unpaired, proximal portion flattened, with a slender distal projection. Gonarcus (fused gonocoxites 11) generally shield-like, sometimes with projections from its posterior surface. Endophallus short, mostly with some bristles or bristle tufts. Female sternum 7 broad, posteriorly produced more or less, with posterior margin truncate or concave; subgenital plate (fused gonocoxites 8) reduced into a small ovoid sclerite or much more modified into a diversely shaped plate. Distribution. Europe (Austria, Czech Republic, European part of Russia, Finland, Germany, Netherlands, Poland, Sweden); Asia (Afghanistan, Armenia, Bhutan, China, India, Japan, Mongolia, North Korea, Pakistan, Russia Far East, Thailand).

Inocellia rara sp. nov. (Figs. 1–10)

Description. Male. Forewing length 7.8 mm. Head (Fig. 1) subquadrate, black, vertex medially with a pair of yellowish brown longitudinal vittae; clypeus reddish brown. Antennal sclerite (torulus) yellowish brown; antennae brown except for scape, pedicel and proximal four flagellomeres which are yellowish brown. Mouthparts reddish brown. Thorax (Fig. 1) black; meso- and metanotum each with a yellowish brown marking medially anterior to yellowish brown scutellum. Legs yellowish brown, slightly darker on tibiae, with yellowish setae. Wings hyaline, pterostigma pale brown, veins brown. Rs with one forked vein and one simple vein running to wing margin. Abdomen (Fig. 1) blackish brown; each pregenital segment with yellowish transverse marking on posterior margin of both tergite and sternite; genital segments brown, but tergum 9 much darker and ectoproct yellow. Tergum 9 (Fig. 2) approximately twice as long as sternum 9, anterior margin slightly incised medially, posterior margin shallowly incised. Sternum 9 (Fig. 4) arcuate, with anterior margin prominent. Gonocoxite 9 (Figs 2–5) shell-like, moderately longer than width of its proximal portion, with apex rounded in lateral view; stylus (gonostylus 9) present subdistad on inner side of gonocoxite 9, short and subtriangular with apex rounded and covered by an inconspicuous dentation, proximally slightly protruding dorsad, which is visible only in lateral view, and digitiform, obliquely directed anteriad in both dorsal and ventral view. Pseudostyli (gonapophyses 9) (Figs 4– 6) feebly sclerotized, foliate, slightly curved anteriorly in ventral view. Fused parameres (complex of gonocoxites, gonapophyses, gonostyli 10) (Figs 5–6, 8) flattened on proximal portion with marginal area curved posteriorly, ventrally bearing a slender and long distal projection, which is slightly curved dorsad and bifurcated at tip. Gonarcus (fused gonocoxites 11) (Figs 4–7) broad, subtrapezoidal and arcuately concaved on ventral margin in caudal view, dorsally with a distally bifurcated projection, and ventrally with a pair of small denticulate processes. Endophallus (Figs 4–6) short, lateral portions distally with a pair of bristle tufts directed anterodorsad and proximally with a pair of bristle tufts directed anteroventrad. Scabrous membranous, narrowly foliate serratulum present (Figs 4–6). Ectoproct (Figs 2–3) subquadrate in lateral view. Hypandrium internum (Figs 6, 9) small with lateral lobes foliate and slightly sinuated posteriorly. Type material. Holotype ♂, CHINA: “North Formosa [= Taiwan], Mt. Taihei [= Mt. Taipingshan in Yilan County, 24°29′50.26″N, 121°32′2.23″E], 2300 m, 28.VI.1941, Coll. T. Sato” (NIAES). Etymology. The specific epithet “rara” refers to the rareness of this new species. It is an adjective in the nominative feminine singular.

Distribution. (Fig. 10) This new species is currently known only from its type locality, i.e. Mt. Taipingshan, in northern Taiwan, China. Remarks. The new species clearly belongs to the I. fulvostigmata species group due to the male gonocoxite 9 which is wider than long and the presence of a scabrous membranous serratulum (see identification key to species of Inocellia in Liu et al. 2012b). The new species appears to be closely related to Inocellia sinensis Navás in having similar male genitalia with the gonocoxite lacking inner bristle tuft, the fused parameres bearing long distal projection, and the double protruding gonarcus. Inocellia rara sp. nov. can be distinguished from I. sinensis by the male gonocoxite 9 with a subtriangular stylus, which is rounded and covered by a number of tiny dentations distally, and by the gonarcus with a pair of projections near dorsal margin and with arcuately concaved ventral margin. In I. sinensis, the stylus of male gonocoxite 9 is unguiform, without any dentations, and the gonarcus has a pair of projections at middle but lacks distinctly concaved ventral margin.

Discussion

The discovery of a third species of Inocelliidae in Taiwan is a great surprise and in particular also with respect to the fact that the new species belongs to the fulvostigmata group of the genus Inocellia. The other two species so far known only from Taiwan can clearly be attributed to the crassicornis group of the genus Inocellia. The species of

FIGURES 2–9. Inocellia rara sp. nov., holotype male. 2—genital segments, lateral view; 3—genital segments, dorsal view; 4—genital segments, ventral view; 5—genital segments, caudal view; 6—ninth gonocoxite with internal structures, lateral view; 7—gonarcus, dorsal view; 8—fused parameres, dorsal view; 9—hypandrium internum, ventral view. Abbreviations: e— ectoproct; ep—endophallus; g—gonarcus (= fused gonocoxites 11); gx9—gonocoxite 9; h—hypandrium internum; p—fused parameres (complex of fused gonocoxites, gonapophyses, and gonostyli 10); ps—pseudostyli (= gonapophyses 9); s— gonostylus 9; S—sternum; ser—serratulum; T—tergum.

230 · Zootaxa 3753 (3) © 2014 Magnolia Press LIU ET AL. The species of the fulvostigmata group have so far only been found in the mainland of Asia, namely in Afghanistan, Pakistan, India (Kashmir), Bhutan, northern Thailand, and eastern parts of China: Inocellia fulvostigmata fulvostigmata U. Aspöck & H. Aspöck, 1968 (Afghanistan, Pakistan, Kashmir), I. fulvostigmata nigrostigmata H. Aspöck, U. Aspöck & Rausch, 1982 (Kashmir), I. bhutana H. Aspöck, U. Aspöck & Rausch, 1991 (Bhutan), I. bilobata U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (northern Thailand), I. cornuta U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (northern Thailand), I. longispina U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (northern Thailand), I. striata U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (northern Thailand), I. cheni Liu, H. Aspöck, Yang & U. Aspöck, 2010 (southern China), I. hainanica Liu, H. Aspöck, Bi & U. Aspöck, 2013 (southern China), I. hamata Liu, H. Aspöck, Yang & U. Aspöck, 2010 (eastern China), I. obtusangularis Liu, H. Aspöck, Yang & U. Aspöck, 2010 (southwestern China), I. sinensis Navás, 1936 (eastern China). Of these I. sinensis is the most closely related species to I. rara sp. nov. by sharing more similar characters than with other Inocellia species, e.g. the similar male genitalia with the gonocoxite lacking inner bristle tuft, the fused parameres bearing long distal projection, and the double protruding gonarcus. Most probably these two species form the sister group to I. hamata. So far, all six species of Raphidioptera known from Taiwan – four species of Mongoloraphidia (Raphidiidae) and two species of Inocellia—are apparently endemic to this island, and most probably also the new species, I. rara, is endemic to Taiwan. This is surprising since throughout the whole glacial period there were frequent broad connections between Taiwan and the Asian mainland. During the Pleistocene the sea level dropped repeatedly by 130 m (possibly even 140 m) compared to the present sea level. This led to an extension of the coast lines of the east of Asia by 600 to 1000 km eastwards. Such events occurred at least one million years, 500 000 years, and 300 000 years B.P., and also during the last glacial period the sea level dropped by 120 m during the maximum of glaciation (ca. 21 000 years B.P.), which led to a broad connection of Taiwan with the mainland of China (Smith et al. 1995, Zhuo et al. 1998, Keally 2005, Liu et al. 2010a). With respect to the marked morphological differences in the male genitalia it is likely that the three species of Inocelliidae (and also the four species of Raphidiidae) of Taiwan are to be traced back to early immigrations, possibly even in the early Pleistocene. The strong orographic structure of the island with its high isolated mountains was undoubtedly a significant precondition for the genetic isolation of the invaders with subsequent endemism. Climate changes resulted in shifts of vertical distributions and enabled species to survive during unfavorable periods due to the possibility of inhabiting optimal altitudes. We must, however, admit that the Raphidioptera fauna of the mountains opposite of Taiwan on the mainland of China is still almost unknown so that definite biogeographical conclusions must await sufficient exploration of these regions of the mainland of China.

Acknowledgements

We would like to thank Dr. Yukinobu Nakatani (Tsukuba) for loan of the specimens from NIAES. We are also grateful to Dr. Masaaki Tomokuni (Tsukuba) for kindly arranging the trip of the first author to the collection of NIAES. We thank Dr. John Plant (Vienna) for linguistic improvement. The research was supported by the National Natural Science Foundation of China (No. 31322501 and 41271063) and the Foundation for the Author of National Excellent Doctoral Dissertation of PR China (No. 201178).

References

Aspöck, H. & Aspöck, U. (1985) Inocellia taiwana n. sp. — eine neue Inocelliiden-Spezies aus Taiwan (Neuropteroidea: Raphidioptera: Inocelliidae). Entomologische Zeitschrift mit Insektenbörse, 95, 45–48. Aspöck, H. & Aspöck, U. (2013) Woher kommen die Namen? Die validen rezenten Taxa der Kamelhalsfliegen der Erde: Systematisches Verzeichnis und Etymologie (Insecta: Endopterygota: Neuropterida: Raphidioptera). Entomologica Austriaca, 20, 9–155. Aspöck, H., Aspöck, U. & Rausch, H. (1991) Die Raphidiopteren der Erde. Eine monographische Darstellung der Systematik, Taxonomie, Biologie, Ökologie und Chorologie der rezenten Raphidiopteren der Erde, mit einer zusammenfassenden Übersicht der fossilen Raphidiopteren (Insecta: Neuropteroidea). 2 Vols. Goecke & Evers, Krefeld, 730 pp & 550 pp. Aspöck, H., Liu, X.Y. & Aspöck, U. (2012) The family Inocelliidae (Neuropterida: Raphidioptera): A review of present knowledge. Mitteilungen der Deutschen Gesellschaft für allgemeine und angewandte Entomologie, 18, 565–573.

NEW INOCELLIA SPECIES FROM TAIWAN Zootaxa 3753 (3) © 2014 Magnolia Press · 231 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 Aspöck, U., Liu, X.Y. & Aspöck, H. (2009) Inocellia shinohara n. sp. — Überraschender Nachweis einer zweiten Spezies der Familie Inocelliidae in Taiwan (Raphidioptera). Entomologische Nachrichten und Berichte, 53, 115–120. Aspöck, U., Liu, X.Y., Rausch, H. & Aspöck, H. (2011) The Inocelliidae of Southeast Asia: A review of present knowledge (Raphidioptera). Deutsche Entomologische Zeitschrift, 58, 259–274. http://dx.doi.org/10.1002/mmnd.201100029 Keally, C.T. (2005) Japanese Pleistocene Landbridges and the Earliest Watercraft. Japanese Archaeology [homepage on the internet] Tama Cable Network, Tokyo, Japan. Available from: http://www.tnet.ne.jp/~keally/MiddlePalaeol/landbridges.html (accessed 21 July 2010) Liu, X.Y., Aspöck, H., Hayashi, F. & Aspöck, U. (2010a) New species of the snakefly genus Mongoloraphidia (Raphidioptera: Raphidiidae) from Japan and Taiwan, with phylogenetic and biogeographical remarks on the Raphidiidae of Eastern Asia. Entomological Science, 13, 408–416. http://dx.doi.org/10.1111/j.1479-8298.2010.00401.x Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2009a) Discovery of Amurinocellia H. Aspöck & U. Aspöck (Raphidioptera: Inocelliidae) in China, with description of two new species. Zootaxa, 2264, 41–50. Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2009b) Inocellia elegans sp. n. (Raphidioptera, Inocelliidae) — A new and spectacular snakefly from China. Deutsche Entomologische Zeitschrift, 56, 317–321. http://dx.doi.org/10.1002/mmnd.201200018 Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2010b) The Inocellia crassicornis species group (Raphidioptera: Inocelliidae) in mainland China, with description of two new species. Zootaxa, 2529, 40–54. Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2010c) Species of the Inocellia fulvostigmata group (Raphidioptera: Inocelliidae) from China. Deutsche Entomologische Zeitschrift, 57, 223–232. http://dx.doi.org/10.1002/mmnd.201300007 Liu, X.Y., Aspöck, H., Zhan, C.H. & Aspöck, U. (2012a) A review of the snakefly genus Sininocellia (Raphidioptera, Inocelliidae): discovery of the first male and description of a new species from China. Deutsche Entomologische Zeitschrift, 59, 235–243. http://dx.doi.org/10.1002/mmnd.200900029 Liu, X.Y., Aspöck, H., Zhang, W.W. & Aspöck, U. (2012b) New species of the snakefly genus Inocellia Schneider, 1843 (Raphidioptera: Inocelliidae) from Yunnan, China. Zootaxa, 3298, 43–52. Liu, X.Y., Aspöck, H., Bi, W.X. & Aspöck, U. (2013) Discovery of Raphidioptera (Insecta: Neuropterida) in Hainan Island, China, with description of a new species of the genus Inocellia Schneider. Deutsche Entomologische Zeitschrift, 60, 59– 64. http://dx.doi.org/10.1002/mmnd.201000019 Smith, A.G., Smith, D.G. & Funnel, B.M. (1995) Atlas of Mesozoic and Cenozoic Coastlines. Cambridge University Press, Cambridge. Zhuo, Z., Baoyin, Y. & Petit-Maire, N. (1998) Paleoenvironments in China during the Last Glacial Maximum and the Holocene Optimum. Episodes, 21, 152–158.