(Raphidioptera: Inocelliidae) from Yunnan, China

Zootaxa 3298: 43–52 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

New species of the snakefly genus Inocellia Schneider, 1843 (Raphidioptera: Inocelliidae) from Yunnan, China XINGYUE LIU1,6, HORST ASPÖCK2, WEIWEI ZHANG3 & ULRIKE ASPÖCK4,5 1

Department of Entomology, China Agricultural University, Beijing 100193, China. Email: [email protected] Department of Medical Parasitology, Institute of Specific Prophylaxis and Tropical Medicine, Medical University of Vienna, Kinderspitalgasse 15, A-1095 Vienna, Austria. Email: [email protected] 3 P.O.Box 4680, Chongqing 400015, China. Email: [email protected] 4 Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, A-1010 Vienna, Austria. Email: [email protected] 5 Department of Evolutionary Biology, University of Vienna, Althanstra嘽e 14, A-1090 Vienna, Austria. Email: [email protected] 6 Corresponding author. E-mail: [email protected] 2

Abstract Two new species of the snakefly genus Inocellia Schneider, 1843 from Yunnan Province of China are described: Inocellia nigra sp. nov. and Inocellia yunnanica sp. nov. A revised key is provided for the males of all Inocellia species. The biogeographical significance of the Yunnanian refugial centre is discussed. Key words: snakefly, Inocellia, Yunnan, China

Introduction Yunnan, an inland province at a low latitude and high elevation, lying between 21°09′–29°15′N and 97°32′– 106°12′E in southwestern China, occupies a vast territory with a diversified and unique fauna and flora. At present, about 12,000 species of insects have been recorded from Yunnan, which represents about 23.5% of the total number of insect species presently known from China (Yang et al. 2004). Due to the presence of three geographic regions (the eastern Asia monsoon region, the Tibetan plateau region and the tropical monsoon region of southern Asia and Indo-China) and the great elevation differences, Yunnan province possesses an extraordinarily rich diversity of species and ecosystems. The fauna of Neuropterida in Yunnan is very abundant, with 15 families and more than 200 species recorded (Yang 1986; Yang & Liu 2010). However, there is only one described species of Raphidioptera in Yunnan, namely Inocellia cheni Liu, H. Aspöck, Yang & U. Aspöck, 2010. Considering the rich fauna of the family Inocelliidae in the Oriental realm (H. Aspöck et al. 1991; U. Aspöck et al. 2011; Liu et al. 2009a, 2009b, 2010a, 2010b), more species of Inocelliidae should be expected in Yunnan. In this paper, we describe two new species of the genus Inocellia Schneider, 1843 based on newly obtained or collected materials from Yunnan. An updated key to the males of all species of Inocellia Schneider, 1843 is given. The significance of the Yunnanian refugial centre for Raphidioptera is discussed in a biogeographical context.

Material and methods Specimens for the present study are deposited in the Entomological Museum of China Agricultural University, Beijing (CAU) and the H. & U. Aspöck collection, Vienna (HUAC). 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).

Accepted by S.L. Winterton: 12 Apr. 2012; published: 3 May 2012

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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 invisible. 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 subapical 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). CommentsThe heuristic value, as well as the conflicts of subdividing the genus Inocellia into the I. crassicornis and I. fulvostigmata groups have been discussed elsewhere (Liu et al. 2010a, 2010b, U. Aspöck et al. 2011).

Key to males of Inocellia 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

Wing membrane hyaline (except pterostigma), immaculate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Wing membrane distinctly brown apically . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. elegans Liu, H. Aspöck, Yang & U. Aspöck Gonocoxite 9 wider than long (Fig. 8)………….(I. crassicornis group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Gonocoxite 9 more or less produced posteriad, slightly or distinctly longer than wide (Fig. 12; Liu et al. 2010b: figs. 17, 28; U. Aspöck et al. 2011: fig. 45)………….(I. fulvostigmata group) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 Gonarcus (fused gonocoxites 11) with paired or unpaired protrusions posteriad (Liu et al. 2010a: fig. 31) . . . . . . . . . . . . . . . 4 Gonarcus without posterior protrusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Posterior protrusion of gonarcus bifurcate (Liu et al. 2010a: fig. 32) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Posterior protrusion of gonarcus paired, slender and digitiform (Liu et al. 2010a: fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. digitiformis Liu, H. Aspöck, Yang & U. Aspöck Gonocoxite 9 with a feebly prominent stylus on inner side (Liu et al. 2010a: fig. 30) . . . . . . . . . . . . . . . . . . . . . I. fujiana Yang Gonocoxite 9 with a long and acutely pointed stylus on inner side (U. Aspöck et al. 2009: fig. 5). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. shinohara U. Aspöck, Liu & H. Aspöck Endophallus proximally with a pair of processes dorsolaterally (Liu et al. 2010a: fig. 6, see arrow) . . . . . . . . . . . . . . . . . . . . . 7 Endophallus proximally without process (Liu et al. 2010a: fig. 15). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Endophallus distally with a pair of tufts of bristles (H. Aspöck et al. 1991: fig. 2499) . . . . . . . . . . . . . . . . I. japonica Okamoto Endophallus distally with only two bristles or none (Liu et al. 2010a: fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. biprocessus Liu, H. Aspöck, Yang & U. Aspöck Gonocoxite 9 with rows of bundles anterior to stylus (H. Aspöck et al. 1991: fig. 2502) . . .I. taiwana H. Aspöck & U. Aspöck Gonocoxite 9 without tuft anterior to stylus, but sometimes with a rather small process posterior to stylus. . . . . . . . . . . . . . . . 9 Gonarcus subtrapezoidal in caudal view (Liu et al. 2010a: fig. 17); endophallus with two pairs of bristle tufts (Liu et al. 2010a: fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. crassicornis (Schummel) Gonarcus subquadrate in caudal view (Fig. 9); endophallus with only one pair of tufts on dorsal surface (Fig. 9). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. nigra sp. nov. Gonocoxite 9 with digitiform or unguiform stylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Gonocoxite 9 with rather small or broadly inflated and roundly tapered stylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Gonocoxite 9 in caudal view with a rounded lobe ventrally (Fig. 15); gonarcus medially domed, but without conspicuous protrusion posteriad (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Gonocoxite 9 in caudal view without ventral lobe (Liu et al. 2010b: fig. 20); gonarcus medially with conspicuous protrusion posteriad (Liu et al. 2010b: fig. 22) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Endophallus without bristles (H. Aspöck et al. 1991: fig. 2518) . . . . . . . . . . . . . . .I. bhutana H. Aspöck, U. Aspöck & Rausch

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13. 14. 15. 16. 17. 18. 19. 20. -

Endophallus with several bristles and tufts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 Gonocoxite 9 with a slender digitiform stylus (U. Aspöck et al. 2011: fig. 6); endophallus with only one pair of tufts on dorsal surfaces (U. Aspöck et al. 2011: fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. longispina U. Aspöck, Liu, Rausch & H. Aspöck Gonocoxite 9 with an unguiform stylus (Fig. 15); endophallus with five tufts on both dorsal and ventral surface (Figs. 14, 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. yunnanica sp. nov. Gonocoxite 9 with an additional process anterior to stylus (U. Aspöck et al. 2011: fig. 47); stylus thickly digitate, bearing several bristles (U. Aspöck et al. 2011: fig. 47) . . . . . . . . . . . . . . . . . . . . . . . . . . . I. cornuta U. Aspöck, Liu, Rausch & H. Aspöck Gonocoxite 9 without additional process anterior to stylus; stylus unguiform, without bristles . . . . . . . . . . . . . . . . . . . . . . . . 15 Gonocoxite 9 with a tuft anterior to stylus on inner side (Liu et al. 2010b: fig. 21); stylus with a geniculate claw, curved dorsad (Liu et al. 2010b: fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. hamata Liu, H. Aspöck, Yang & U. Aspöck Gonocoxite 9 without tuft anterior to stylus on inner side (Liu et al. 2010b: fig. 39); stylus simply unguiform, slightly curved ventrad (Liu et al. 2010b: fig. 38) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. sinensis Navás Gonocoxite 9 with stylus inconspicuous, rather small, and bearing no bristle or tuft on inner side (Liu et al. 2010b: fig. 7). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. cheni Liu, H. Aspöck, Yang & U. Aspöck Gonocoxite 9 with prominent stylus, broadly inflated, and bearing a tuft on inner side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Gonocoxite 9 with stylus bilobed (U. Aspöck et al. 2011: fig. 13). . . . . . . . . .I. bilobata U. Aspöck, Liu, Rausch & H. Aspöck Gonocoxite 9 with stylus not bilobed.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Gonarcus medially without protrusion (Liu et al. 2010b: fig. 32) . . . . . . I. obtusangularis Liu, H. Aspöck, Yang & U. Aspöck Gonarcus medially with conspicuous protrusions. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Stylus of gonocoxite 9 nearly cone-like (U. Aspöck et al. 2011: fig. 32); gonarcus medially with a distally bifurcated protrusion and with ventral corners acutely angulated (U. Aspöck et al. 2011: fig. 35) . . . I. striata U. Aspöck, Liu, Rausch & H. Aspöck Stylus of gonocoxite 9 lobe-like; gonarcus medially with a pair of protrusions and with ventral corners rounded (H. Aspöck et al. 1991: figs. 2508, 2515, 2516)……….I. fulvostigmata U. Aspöck & H. Aspöck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Pterostigma yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I. fulvostigmata fulvostigmata U. Aspöck & H. Aspöck Pterostigma brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .I. fulvostigmata nigrostigmata H. Aspöck, U. Aspöck & Rausch

Inocellia nigra sp. nov. (Figs. 1–2, 8–11) Diagnosis. The appearance of the species is conspicuously characterized by black elements including head, thorax, wing venation, legs, and abdomen. The male is characterized in the genitalia by the short gonocoxite 9 with a rather short and cone-like stylus, the sub-quadrate gonarcus (fused gonocoxites 11) with ventromedial protrusion, and the endophallus with only one pair of tufts with bristles on dorsal surface. Description. Male. Body length 7.7–8.3 mm; forewing length 6.7–7.7 mm, hindwing length 5.6–6.6 mm. Head (Fig. 1) sub-quadrate, black throughout. Antennal sclerite (torulus) and antennae blackish brown. Mouthparts black, mandibles with distal half reddish-brown. Thorax black throughout. Legs (Fig. 2) blackish brown with brownish setae; coxae, trochanters, and bases of femora yellow. Wings (Fig. 1) hyaline, pterostigma blackish brown, veins blackish brown. Anterior branch of Rs with one forked vein and one simple vein running to wing margin. Abdomen blackish brown; tergum of each pregenital segment medially with a small yellow spot, sternum of each pregenital segment with posterior yellow margin. Genital segments entirely blackish brown. Gonocoxite 9 (Figs. 8–10) dome-like, nearly semicircular, much wider than long; distal portion of inner side with a rather short stylus, median portion of the stylus slightly prominent dorsally, with numerous bristles. Pseudostylus (basal part of gonapophysis 9) (Fig. 9) paired and foliate. Fused parameres (complex of fused gonocoxites, gonapophyses, and gonostyli 10) (Figs. 9–11) small, proximal portion flattened, expanded, subtrapezoidal and concave ventrad, distal projection curved dorsad. Gonarcus (fused gonocoxites 11) (Figs. 9–11) shield-like, with dorsal margin feebly produced posteriorly; in caudal view subquadrate, with ventromedial portion produced into a subtriangular lobe. Endophallus (Figs. 9–10) short, distal portion medially with a pair of bristle tufts directed dorsad. Ectoproct (Fig. 8) subtrapezoidal with rounded posterodorsal corner in lateral view. Hypandrium internum small, lobes narrow. Female. Unknown. Type material. Holotype ♂, CHINA: “Yunnan, Baoshan, along the road from Wayao to Laoyingshan [ca. 25°26′N, 99°15′E, 1800 m], 12.V.2009, Liang Ding” (CAU). Paratype 1♂, CHINA: “Yunnan, Gaoligong mts., 25.22N, 98.49E, 1500–2500 m, 17-24.V. 1995, Vít Kubáň (HUAC). Distribution. The new species is currently known only from the southern part of the Gaoligongshan Range. Etymology. The specific epithet ‘nigra’ refers to the predominately blackish coloration including head, thorax, wing venation, legs and abdomen. It is an adjective in the feminine nominative case from Latin niger, -a, -um = black. NEW INOCELLIA FROM YUNNAN

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FIGURES 1–4. Habitus images of Inocellia spp. 1. I. nigra sp. nov., holotype male; 2. Same, lateral view; 3. I. yunnanica sp. nov., holotype male; 4. Same, paratype female. Scale bars = 1.0 mm.

Remarks. The new species belongs to the Inocellia crassicornis group and appears to be closely related to I. crassicornis by having a tiny stylus near the posterior margin of the inner side of male gonocoxite 9, however, it is distinguished from I. crassicornis in appearance by the dark legs. In I. crassicornis as well as the other Inocellia species, the legs are generally yellow. Considering the male genitalia, the new species differs from I. crassicornis by the subquadrate gonarcus in caudal view and the endophallus with only one pair of tufts. In I. crassicornis the gonarcus is subtrapezoidal in caudal view, and the endophallus has two pairs of tufts of bristles on both dorsal and ventral surfaces.

Inocellia yunnanica sp. nov. (Figs. 3–7, 12–24) Diagnosis. In appearance, the species is entirely black on head and thorax, and dorsally blackish brown with transverse yellowish bands on pregenital segments of the abdomen. The male is characterized in the genitalia by the elongated gonocoxite 9 with a conspicuous unguiform stylus and a subtriangular lobe posteroventrally to stylus. The female is characterized by the tergum 8 posteroventrally with a short rounded protrusion. Description. Male. Body length 10.0 mm; forewing length 8.2 mm, hindwing length 6.6 mm. Head (Fig. 3) subquadrate, black throughout. Antennal sclerite (torulus) and antennae blackish brown. Mouthparts black, mandibles with distal half reddish brown. Thorax black throughout. Legs yellow with yellowish setae. Wings (Fig. 3) hyaline, pterostigma blackish brown, veins blackish brown. Anterior branch of Rs with one forked vein and one simple vein running to wing margin.


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FIGURES 5–7. Inocellia yunnanica sp. nov. 5. Male pupa; 6. Female adult; 7. Collecting site of type specimens in Mt. Wuliangshan.

Abdomen blackish brown with venter much paler; joints of each pregenital segment transversely yellow. Genital segments entirely yellow. Tergum 9 (Figs. 12–13) approximately 1.5 times as long as sternum, anterior margin feebly incised medially, posterior margin nearly truncate. Sternum 9 (Figs. 12–14) arcuate, with anterior margin slightly prominent. Gonocoxite 9 (Figs. 12–16) broadly dome-like, much longer than width of its proximal portion, with apex acutely pointed in lateral view; stylus (gonostylus 9) produced from medial portion of inner side, unguiform and feebly curved ventrad; a rounded lobe posteroventral to the stylus; a bristle tuft anterior to stylus. Pseudostylus (basal part of gonapophysis 9) (Figs. 14–15) paired, feebly sclerotized, narrowly foliate. Fused parameres (complex of fused gonocoxites, gonapophyses, gonostyli 10) (Figs. 16, 18) flattened and subtrapezoidal on proximal portion, ventrally bearing a slender distal projection, which is slightly curved dorsad. Gonarcus (fused gonocoxites 11) (Figs. 15–17) shield-like, nearly trapezoidal in caudal view, medially prominent posteriad. Endophallus (Figs. 14–16) short, dorsally with a pair of short bristle tufts on proximal portion and a pair of long bristle tufts on feebly sclerotized areas of distal portion, ventrally with a short bristle tuft medially. A scabrous membranous structure is present near the proximodorsal portion of endophallus. Ectoproct (Fig. 12) in lateral view subquadrate. Hypandrium internum (Fig. 19) small with lateral lobes foliate and slightly sinuate posteriorly. Female. Forewing length 13.3 mm, hindwing length 11.5 mm. Sternum 7 (Figs. 20–21, 24) nearly trapezoidal in lateral view, posterior margin slightly produced. Tergum 8 (Figs. 20, 22, 24) posteroventrally with a short and round protrusion. Subgenital plate (fused gonocoxites 8) (Figs. 20–21, 24) separated into a broadly subquadrate sclerite anteriorly and a small ovoid sclerite posteriorly, which are connected by a grooved membrane. Atrium bursae (Figs. 20, 23) subtriangular, slightly sclerotized ventrad; sacculus bursae and receptaculum seminis damaged in female paratype, but well preserved in the female specimen from Mt. Meili (Fig. 23).

NEW INOCELLIA FROM YUNNAN


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Type material. Holotype ♂, “CHINA / Yunnan, Puer city, Jingdong county, Wuliangshan Nature Reserve, Gibbon Observation Station [1787 m], N 24°21′07″/E 100°42′12″, April, 2.2010, ZHANG Weiwei leg.” (CAU). Paratype 1♀, same data as holotype (CAU).

FIGURES 8–11. Inocellia nigra sp. nov. 8. Male genital segments, lateral view; 9. Male genital segments, caudal view; 10. Male genital segments showing internal structures, lateral view; 11. Fused parameres (complex of fused gonocoxites, gonapophyses, and gonostyli 10), dorsal view. e: ectoproct; ep: endophallus; g: gonarcus (fused gonocoxites 11); gx9: gonocoxite 9; p: fused parameres; ps: pseudostylus (basal parts of gonapophyses 9); s: stylus or stylus-like process (gonostylus 9); S8, 9: sternite 8 and 9; T8, 9: tergite 8 and 9. Scale bar = 0.5 mm.


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FIGURES 12–19. Inocellia yunnanica sp. nov. 12. Male genital segments, lateral view; 13. Male genital segments, dorsal view; 14. Male genital segments, ventral view; 15. Male genital segments, caudal view; 16. Male genital segments showing internal structures, lateral view; 17. Gonarcus (fused gonocoxites 11), dorsal view; 18. Fused parameres (complex of fused gonocoxites, gonapophyses, and gonostyli 10), dorsal view; 19. Hypandrium internum, ventral view. Scale bar = 0.5 mm.

Other material. 1♀, CHINA: “YUNNAN, 5.-8.vii.1996, 28°06′N 98°54′E, 2700 m, Hengduan mts-part MEILI, Vít Kubáň leg.” (HUAC); 1♀, CHINA: Yunnan, Tengchong, Datang [25°33′N, 98°53′E, 1118 m] (CAU). Distribution. The new species is found to be distributed in three mountain ranges, i.e. Wuliangshan Range in central Yunnan, Gaoligongshan Range in western Yunnan, and Meili in northwestern Yunnan, all of which belong to the Hengduan orographic uplift. Etymology. The specific epithet ‘yunnanica’ refers to the endemic distribution in Yunnan Province. It is an adjective in the feminine nominative case from Latin Yunnanicus, -a, -um = Yunnanian. Remarks. The new species belongs to the Inocellia fulvostigmata group and appears to be closely related to Inocellia longispina from Thailand by having similar male gonocoxite 9 with a conspicuous stylus and a subtriangular lobe. However, the new species differs from I. longispina by the elongated male gonocoxite 9, the unguiform stylus, and the endophallus with five tufts, while in I. longispina the male gonocoxite 9 is much shorter with digitiform stylus and the endophallus possesses only one pair of tufts on the dorsal surface. The female of the new species can be easily distinguished from the other Inocellia species by the tergum 8 with a posteroventral protrusion. Since we did not find male specimens of the new species from the Gaoligongshan and Meili ranges, the current identification of the two female specimens from these two localities was made tentatively; these specimens are not placed in the type series. The following biological notes pertain to the type specimens. These specimens were collected from a naturally dead tree (species unidentified) at the top of a mountain, where some large trees were cut down, leaving a relatively open area. The male was collected as a pupa sitting in a small hole when removing the tree bark in the afternoon (ca. 4:00 pm). The female was collected as a freshly emerged adult on the bark of the same tree in the evening (ca. 9:00 pm).



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FIGURES 20–24. Inocellia yunnanica sp. nov. 20. Female genital segments, lateral view, paratype female; 21. Female genital segments, ventral view, paratype female; 22. Female genital segments, lateral view, female from Mt. Meili; 23. Bursa copulatrix, lateral view, female from Mt. Meili; 24. Female genital segments, lateral view, female from Mt. Gaoligongshan. ab: atrium bursae; gr: glandula receptaculi; o: ovipositor; r: receptaculum seminis; sb: sacculus bursae; sg: subgenitale (fused gonocoxites 8); S7: sternite 7; T7–9: tergite 7–9. Scale bar = 0.5 mm.

Discussion Until a few years ago it was assumed that the east of Asia (where the southern boundary of the occurrence of Raphidioptera lies) harbors only one or two species of snakeflies (H. Aspöck et al. 1991). In recent years this view has been entirely changed. Surprisingly, several new species belonging to the family Inocelliidae had been discovered in the south of China and in the north of Thailand, respectively (Liu et al. 2010a, b; U. Aspöck et al. 2011). Altogether nine species of Raphidioptera, all belonging to the family Inocelliidae, can be attributed to the Yunnanian centre: Inocellia digitiformis Liu, H. Aspöck, Yang & U. Aspöck, 2010 (China: Sichuan) Inocellia nigra sp. nov. (China: Yunnan) Inocellia cheni Liu, H. Aspöck, Yang & U. Aspöck, 2010 (China: Guanxi, Yunnan) Inocellia longispina U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (NW-Thailand) Inocellia bilobata U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (NW-Thailand) Inocellia striata U. Aspöck, Liu, Rausch & H. Aspöck , 2011 (NW-Thailand) Inocellia cornuta U. Aspöck, Liu, Rausch & H. Aspöck, 2011 (NW-Thailand) Inocellia yunnanica sp. nov. (China: Yunnan) Parainocellia burmana (U. Aspöck & H. Aspöck, 1968) (Myanmar) As can be seen from the list, three species have been recorded within Yunnan, one species has been found in the adjacent Province of Sichuan, one species has been described from Myanmar, and four species are known from the northwest of Thailand. There is no other part in the world where such a high number of species occurs in a territory of comparable size. In most parts of the Old World only one species of Inocelliidae occurs (H. Aspöck et al. 2012). One may certainly question whether all these nine species are to be assigned to one single refugial centre, namely


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the Yunnanian centre in the sense of de Lattin (1967) or whether several centres or subcentres within this part of Southeast Asia can be located. There can be no doubt that Thailand lies within the Oriental realm; however, the mountainous regions in the north together with the higher altitudes of Yunnan and Myanmar may be classified as representing a transgression zone between the Palaearctic and the Oriental regions, whereby the percentage of the Oriental elements is apparently the prevailing one. As we have already discussed elsewhere (U. Aspöck et al. 2011), one cannot exclude the possibility that the I. fulvostigmata group of the genus Inocellia has an Oriental origin. Of the three species so far found in Yunnan, one (I. nigra) belongs to the I. crassicornis group (which is certainly of Palaearctic origin), but two (I. cheni, I. yunnanica) belong to the I. fulvostigmata group. Currently, only eight specimens (four males, four females) of Inocelliidae from altogether six localities in Yunnan are known from collections. The fact that they represent three species leads to the assumption that this part of China, which has several hotspots of biodiversity (Yang et al. 2004), may harbor further species.

FIGURE 25. Geographic distribution of the Inocellia species from Yunnan Province. ●: I. nigra sp. nov.; ■: I. yunnanica sp. nov.; : I. cheni Liu, H. Aspöck, Yang & U. Aspöck.

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It is of interest that no specimens of the second family of Raphidioptera, Raphidiidae, have been found in Yunnan, as yet. The nearest records of Raphidiidae are from the Hubei Province. It seems inconceivable that the family Raphidiidae does not occur in Yunnan. Further investigations on snakeflies in Yunnan might lead to additional discoveries.

Acknowledgements We are much indebted to Mr. Liang Ding (Beijing) and Mr. Hao Huang (Shanghai), as well as to Mr. Víteslav Kubáň (Brno) for kindly providing the valuable specimens which were collected in Yunnan. Grateful thanks go to Prof. Dr. Peter Comes (Salzburg) and to Prof. Dr. Thomas Schmitt (Trier) for valuable discussions on biogeography. We thank Dr. John Plant (Vienna) for linguistic improvement. The research was supported by the National Natural Science Foundation of China (Nos. 31000973 and 31110103002) and the Foundation for the Author of National Excellent Doctoral Dissertation of PR China (No. 201178).

References 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, in press. Aspöck, H., Aspöck, U. & Rausch, H. (1991) Die Raphidiopteren der Erde. Eine monographische Darstellung der Systematik, Taxonomie, Biologie, 嘵kologie und Chorologie der rezenten Raphidiopteren der Erde, mit einer zusammenfassenden Übersicht der fossilen Raphidiopteren (Insecta: Neuropteroidea). Goecke & Evers, Krefeld, 2 volumes, 730 pp, 550 pp. Aspöck, U. & Aspöck, H. (2008) Phylogenetic relevance of the genital sclerites of Neuropterida (Insecta: Holometabola). Systematic Entomology, 33, 97–127. 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. Lattin, G. de (1967) Grundri嘽 der Zoogeographie. VEB Gustav Fischer Verlag, Jena, 602 pp. 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. Liu, X.Y., Aspöck, H., Yang, D. & Aspöck, U. (2010a) 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. (2010b) Species of the Inocellia fulvostigmata group (Raphidioptera: Inocelliidae) from China. Deutsche Entomologische Zeitschrift, 57, 223–232. Yang, C.K. (1986) Thirty new species and four new genera of Neuroptera from Yunnan, and the family Nemopteridae new to China. Acta Agriculturae Universitatis Pekinensis, 12, 153–166, 423–434. (in Chinese with English summary) Yang, D. & Liu, X.Y. (2010) Fauna Sinica Insecta Vol.51 Megaloptera. Science Press, Beijing. 457 pp, XIV pl. (in Chinese with English summary) Yang, Y.M., Tian, K., Hao, J.M., Pei, S.J. & Yang, Y.X. (2004) Biodiversity and biodiversity conservation in Yunnan, China. Biodiversity and Conservation, 13, 813–826.


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