Revision of the Fishfly Genus Ctenochauliodes Van Der Weele

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RevisionBlackwell Publishing Ltd of the ﬁshﬂy genus Ctenochauliodes van der Weele (Megaloptera, Corydalidae)

XING YUE LIU & DING YANG

Accepted: 15 April 2006 Liu, X. Y. & Yang, D. (2006). Revision of the fishfly genus Ctenochauliodes van der Weele doi: 10.1111/j.1463-6409.2006.00240.x (Megaloptera, Corydalidae). — Zoologica Scripta, 35, 473–490. The first taxonomic revision and phylogenetic analysis of the Oriental fishfly genus Cteno- chauliodes van der Weele, 1909 is presented. Except for C. punctulatus, where the male is unknown, all species of Ctenochauliodes and two outgroups (Archichauliodes chilensis Kimmins and Protochauli- odes bullocki Flint) were scored for 24 adult morphological characters, 21 of which were obtained from the male genitalia. The monophyly of Ctenochauliodes is corroborated by a series of synapomorphic characters. The pectinate female antenna was proposed as a diagnostic character. Two main clades within Ctenochauliodes were found based on the present phylogenetic analysis. All 12 Ctenochauliodes species are keyed, described and illustrated, with five species new to science. Ctenochauliodes forcipatus Kimmins is a junior synonym of C. friedrichi Navás, and C. moganshanus Yang & Yang is a junior synonym of C. griseus Yang & Yang. All Ctenochauliodes species were recorded from southern China, including 11 endemic species. Hence, there is no doubt that southern China should be acknowledged as the distribution centre of Ctenochauliodes. Xingyue Liu, Department of Entomology, China Agricultural University, Beijing, 100094, China. E-mail: [email protected] Ding Yang, Department of Entomology, China Agricultural University, Beijing, 100094, China; Key Laboratory of Insect Evolution & Environmental Changes, Capital Normal University, Beijing 100037, China. E-mail: [email protected]

Introduction Chauliodes nigrovenosus van der Weele from ‘Tonkin’ (now The megalopteran genus Ctenochauliodes van der Weele is a small northern Vietnam) as the type species. Navás (1932) described genus within the subfamily Chauliodinae. All Ctenochauliodes a second Ctenochauliodes species from Sichuan, China. Later, species occur only in the Oriental Realm and are particularly Kimmins (1954) established a third species, Ctenochauliodes concentrated in China. Due to the low population density in forcipatus, also from Sichuan, China. Following this description, the field and dull appearance, the genus is not distinctly strik- the genus was not recorded or described for the next 30 years. ing in megalopteran collections and not well studied. Since 1986, systematic studies on Ctenochauliodes have been In comparison with the other genera of Chauliodinae, made by Yang & Yang (1986, 1990, 1992a,b, 1999), which Ctenochauliodes is small in size, usually with dispersed brownish resulted in the description of six additional new species. The wing patterns. The remarkable diagnostic character is the centre of distribution for Ctenochaliodes was predicted to be strongly developed pectinate male antenna, whereas the female China (Yang & Yang 1992b). antenna is only moderately developed. The male genitalia of According to a study on the generic phylogeny of Chaulio- Ctenochauliodes possesses highly modified features, characterized dinae (Liu & Yang 2006b), the genus Ctenochauliodes is assigned by the following characters: tenth tergum apically twisted or to a monophyletic clade which comprises all Asian fishfly with one ventral process produced from its subapical portion; genera and two Nearctic genera (Nigronia and Chauliodes). male tenth sternum in lateral view much wider on apical half, Ctenochauliodes is assigned as the stem group to the other genera and bifurcated from its subbasal portion in ventral view, within this clade. However, the monophyly of Ctenochauliodes forming a pair of asymmetric lobes. has not been tested using a dataset that includes all species of Van der Weele (1907) erected the genus Ctenochauliodes for the genus, and its interspecific phylogeny has not been studied species with pectinate antennae in both sexes and designated so far. Therefore, in the present study all Ctenochauliodes

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Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

species were revised, with five species described as new to Appendix 1). The cladogram was rooted using the following science. The monophyly of Ctenochauliodes was retested and two outgroup species, Archichauliodes chilensis Kimmins and the phylogenetic relationships were analysed at the species Protochauliodes bullocki Flint. The outgroup selection was level. based on the generic phylogeny of Chaulodinae (Liu & Yang 2006b). The selected outgroups belong to the clade which Materials and methods was assigned as the sister to the clade which comprised all Taxa and terminology Asian fishfly genera and two Nearctic genera (Nigronia and All specimens examined were obtained from the Entomological Chauliodes). Furthermore, Archichauliodes has been considered Museum of the China Agricultural University (CAU), the most primitive genus in the subfamily Chauliodinae (van Beijing, and the Institute of Zoology, Chinese Academy of der Weele 1910). Science (IZCAS), Beijing. The holotype of Ctenochauliodes The ingroup sampling included all species of Ctenochauliodes forcipatus Kimmins, whose habitus photo was examined, is except C. punctulatus, where the male is unknown. If adding deposited in the Natural History Museum (NHM), London, C. punctulatus, any resolution of this species cannot be obtained UK. The outgroup species from the New World were because most morphological data are unknown. obtained through a kind exchange by Dr O.S. Flint, Jr. The present cladistic analysis was performed using PAUP* from the National Museum of Natural History (NMNH), ver. 4.0b10 (Swofford 2002) employing a heuristic parsimony Washington, DC, USA. analysis, with 1000 random stepwise additions of taxa (TBR The specimens were collected mostly at light traps in branch swapping) under ACCTRAN optimization, characters mountainous regions. Genitalic preparations were made by unordered and of equal weight, MulTrees option in effect. macerating the apex of the abdomen in cold 10% KOH for Bootstrap values for clades were calculated in 1000 replicates 8–10 h. The apex of the abdomen was then transferred to using a general heuristic search, the branches with bootstrap glycerol for further dissection and examination. After exam- values < 50% collapsed. Bremer’s decay index was calculated ination it was moved to fresh glycerol and stored in a microvial with Autodecay ver. 4.0 (Eriksson 1998) and PAUP* ver. 4.0b10. pinned below the specimen. The habitus images of adults were The unambiguous characters were mapped by MacClade ver. taken with a Nikon Coolpix 4500 digital camera and can be seen 4.0 (Maddison & Maddision 2000). in Morphobank (http://www.morphobank.com). Accession numbers in Morphobank are indicated in the headings of Results each species in the systematics section. Drawings were made The heuristic analysis resulted in three most parsimonious under the light microscope. trees (length = 34, consistency index (CI) = 0.82, retention The terminology of the adult genitalia generally follows index (RI) = 0.88). The three MP trees differ only in the Contreras-Ramos (2004). placement of C. digitiformis. This species was either assigned as the sister of C. abnormis + C. friedrichi, the sister of Cladistic analysis C. sagittiformis, or unresolved. One of the three MP trees The phylogenetic analysis includes 24 adult morphological is shown in Fig. 1, with unambiguous characters mapped. characters, of which 21 are male genitalic characters (Table 1; In the strict consensus tree for the three MP trees (Fig. 2),

Table 1 Character state distribution used in cladistic analysis.

Characters 123456789101112131415161718192021222324

Taxa Archichauliodes chilensis 000000000000000000000000 Protochauliodes bullocki 000000000000000000000000 Ctenochauliodes abnormis 1?101000011101100010012? Ctenochauliodes digitiformis 1?110000011110001010001? Ctenochauliodes elongatus 110001111111010001210001 Ctenochauliodes friedrichi 111010000111100000000120 Ctenochauliodes fujianensis 1?000110011101000021100? Ctenochauliodes griseus 110001100111010001211000 Ctenochauliodes meridionalis 110001111111010001210201 Ctenochauliodes nigrovenosus 110001111111010001210200 Ctenochauliodes sagittiformis 1?100000011110010000000? Ctenochauliodes similis 110001111111010000210200 Ctenochauliodes yangi 110001000111010000000202

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X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Fig. 1 One of the three MP trees with Archichauliodes chilensis and Protochauliodes bullocki as the outgroups (see text for further details). Only unambiguous characters are shown. Shaded squares represent apomorphic characters, those with open squares represent parallelisms and reversals. Character states for multistate characters and reversals are placed above the squares.

the ingroup is divided into two monophyletic clades. Clade sparsely borne on the male tenth tergum are shared only by 1 comprises C. abnormis, C. digitiformis, C. friedrichi, and the Ctenochauliodes species, which corroborated the monophyly C. sagittiformis. Within clade 1, the placement of C. digitiformis of the genus. and C. sagittiformis was unresolved forming a polychotomy. The other Ctenochauliodes species formed clade 2. Within clade 2, a Clade 1 monophyletic subclade was formed by C. elongatus, C. fujianensis, The monophyly of clade 1 was supported by the apically C. griseus, C. meridionalis, C. nigrovenosus, and C. similis, while twisted male tenth tergum (char. 3: 1) and the male tenth C. yangi was assigned as the sister to the subclade. sternum which is not curved dorsad (char. 13: 1). Clade 1 was considered to be relatively more primitive than clade 2 Discussion because the four included species possess plesiomorphic Monophyly of Ctenochauliodes characters, such as the simple male tenth tergum without According to the present result, the monophyly of the genus ventral process and the male tenth sternum with both Ctenochauliodes was well supported by a series of synapo- lobes blunt at the tip and of equal length. The sister relation- mophic characters, e.g. the pectinate male antenna (char. ships between C. abnormis and C. friedrichi were well supported 1: 1), the pectinate female antenna (char. 2: 1), the stout by the anteriorly twisted tip of the male tenth tergum (char. spinous setae sparsely borne on the male tenth tergum 5: 1) and the presence of the sclerite beneath the male anus, (char. 10: 1), the deeply bifurcated male tenth sternum which is not trilobed (char. 22: 1). Though the assignment of (char. 11: 1), and the asymmetric lobes of the male tenth sternum C. digitiformis was not conﬁrmed, we presume this species to (char. 12 : 1). Within the synapomorphic characters above, be the sister of the C. abnormis and C. friedrichi subclade by the pectinate female antenna and the stout spinous setae the presence of the lateral lobes of the male tenth sternum.

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Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

Fig. 2 The strict consensus tree of three most parsimonious trees with bootstrap values (above branches) and Bremer’s decay indices (below branches).

Clade 2 The monophyletic subclade of C. elongatus + C. fujianensis The monophyly of clade 2 was supported by the presence of + C. griseus + C. meridionalis + C. nigrovenosus + C. similis was the ventral process on the male tenth tergum (char. 6: 1) and well supported by the well developed ventral process on the the acutely pointed left lobe of the male tenth sternum (char. male tenth tergum (char. 7: 1), the acutely pointed right lobe 14: 1). According to its appearance, C. yangi is somewhat of the male tenth sternum (char. 19: 2), and short rounded related to C. abnormis in having the similar striking wing lateral arms of the male tenth sternum (char. 20: 1). Cteno- pattern. However, in the present study this species is the chauliodes elongatus, C. meridionalis, C. nigrovenosus, and C. similis sister group to the subclade (C. elongatus + C. fujianensis + were grouped by the male tenth tergum with an acutely pointed C. griseus + C. meridionalis + C. nigrovenosus + C. similis). tip (char. 9: 1) and the presence of subtriangular ventral Hence, the striking wing pattern may be a parallel derived processes (char. 8: 2). The male tenth sternum with the right lobe character for C. abnormis and C. yangi. The basal status of longer than the left (char. 18: 1) weakly supported the group- C. yangi was obtained by a series of plesiomorphic characters ing of C. elongatus, C. meridionalis, and C. nigrovenosus. The present in this species, such as the male tenth tergum with a acutely pointed female gonocoxite (char. 24: 1) is synapomorphic blunt tip, the feebly developed ventral process of the male for the C. elongatus and C. meridionalis sister pairing. Cteno- tenth tergum, the elongate and subtriangular lateral arms of chauliodes fujianensis and C. griseus were grouped as the sister the male tenth sternum, and the male tenth sternum with the group of the subclade comprising C. elongatus + C. meridionalis left lobe as long as the right lobe. + C. nigrovenosus + C. similis. The monophyly of the former

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

subclade is supported by the male tenth sternum not narrowed behind the lateral arm in lateral view (char. 21: 1).

Zoogeography of the Ctenochauliodes species All the Ctenochauliodes species are known to occur in the Oriental Realm and are especially concentrated in southern China. Eleven Ctenochauliodes species are endemic to China, with C. nigrovenosus also distributed in northern Vietnam. Therefore, we conclude that the distribution centre of Ctenochauliodes is in southern China, as first suggested by Yang & Yang (1992b). Most Ctenochauliodes species are restricted in distribution and sympatric patterns are rarely presented. Under the system of ‘Zoogeographical Regions of China’ erected by Zhang (1998), three major regions (South-west China, Fig. 4 Map showing the geographical distributions of Ctenochauliodes Central China, and South China) are recognized as the elongatus (), C. fujianensis (), C. griseus (), C. meridionalis (), distribution areas for the species of Ctenochauliodes. Among C. nigrovenosus (), C. similis (), and C. yangi (). Ctenochauliodes, C. friedrichi is most widely distributed, being present in all three regions. Besides C. friedrichi, there are eight species (C. abnormis, C. digitiformis, C. meridionalis, The previous studies of the zoogeography of Megaloptera C. nigrovenosus, C. punctulatus, C. similis, C. sagittiformis, and showed that the current Asian Corydalidae fauna may have C. yangi ) in South China, while there are seven species in originated from north-eastern India or south-western China Central China, with four species (C. elongatus, C. griseus, during the split of Laurasia and Gondwanaland or the C. fujianensis, and C. sagittiformis) endemic to this region. connection of the Indian subcontinent to Eurasia (Penny Considering the distribution patterns mentioned above 1993; Liu et al. 2005). According to the result of the cladistic (Figs 3, 4), a moist tropical climate is obviously preferred by analysis, the centre of origin of Ctenochauliodes might also fit species of Ctenochauliodes because approximately 70% of them with the centre of origin of Asian Corydalidae mentioned are distributed in the South China Region. It is inferred from previously because the assumed primitive species were the narrowly disjunct distribution areas of most Cteno- distributed in south-western China. Compared with other chauliodes species that: (1) their ability to disperse may be very megalopteran genera, the diversity of Ctenochauliodes is much weak; (2) the larvae may be rigorously adapted to extremely lower in the presumed centre of origin, with only two species clean aquatic habitats, and (3) their competitive ability may distributed (Liu et al. 2005; Liu & Yang 2005, 2006a). How- be somewhat poorer than some other Oriental fishfly genera. ever, the populations of Ctenochauliodes may be quite rare and consequently poorly sampled. Therefore, it is reasonable to believe that more species of Ctenochauliodes will be found within the presumed centre of origin. The distribution track of Ctenochauliodes is herein recognized from the centre of origin to the eastern areas. Ctenochauliodes punctulatus is the single species that occurs in the island of Hainan. However, until now, Ctenochauliodes has not been found in Taiwan, although habitats appear to be suitable for this genus. On the basis of the present known distribution patterns, the fauna of Hainan may be more closely related to the fauna of the Chinese mainland than to Taiwan.

Systematics Genus Ctenochauliodes van der Weele, 1909 Ctenochauliodes van der Weele, 1909. Not. Leyden Mus. 30: 263. Type species: Chauliodes nigrovenosus Walker, 1853. Fig. 3 Map showing the geographical distributions of Ctenochauliodes abnormis (), C. digitiformis (), C. friedrichi (), C. punctulatus (), General characters. Fishﬂies of small to medium size (12– and C. sagittiformis (). 23 mm), with yellowish-brown to black body. Wings hyaline

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

or subhyaline, usually with dark markings. Head subtriangular subquadrate and not apically produced; ninth tergum deep and robust with distinctly prominent compound eyes. Antennae and pointed ventrad; tenth tergum short claviform, usually pectinate in both sexes, but more strongly developed in with apical half strongly narrowed, and with strongly pro- male with median branches much longer. Labrum suboval. minent cerci; gonocoxite subtriangular with more or less Prothorax subcylindrical, not longer than wide. Meso- and pointed tips. metathorax distinctly robust. Legs with dense short setae. Wings somewhat narrow, about 3.0 times longer than wide, Natural history. Generally, the larvae are aquatic and the adults

with roundly pointed tips; Rs four-branched, with R2 two to are nocturnal. Label data indicate that adults emerge from four-branched; three crossveins between R1 and Rs; M two- early April to late July. Most species were captured in the branched; 1A two-branched, with posterior branch strongly national forest reserves, showing that these insects prefer sinuated; 2A two-branched, with both branches strongly relatively undisturbed environments. We also inferred from sinuated; basal r-m crossveins strong and re-connected to the peculiar pectinate female antenna that both sexes may M by short branch. Male ninth tergum subquadrate in dorsal release pheromones which can be received by the pectinate view, much shorter than wide, usually with shallowly arched antenna during courtship. basal incision; ninth sternum less sclerotized and nearly as long as ninth tergum, without membranous subtriangular Remarks. The genus Ctenochauliodes is somewhat similar to apical process; tenth tergum short claviform, as long as or the genus Archichauliodes in usually possessing a bifurcated shorter than ninth tergum, basally thick and distinctly nar- male tenth sternum, while the deep and pointed male ninth rowed towards tip, with strongly prominent cercus, and usu- tergum shows that Ctenochauliodes also has close affinities ally with one ventral process somewhat produced inwards; with the Oriental fishfly genera Anachauliodes and Neochauli- male tenth sternum strongly sclerotized, longer than ninth odes. However, it can be easily distinguished from the other tergum, in lateral view with apical half somewhat widened, genera of Chauliodinae by the pectinate antenna in both in ventral view bifurcated from subbasal portion, forming sexes and the male tenth sternum bearing a pair of asymmetric pair of asymmetric lobes. Female eighth sternum mostly lobes.

Key to species of Ctenochauliodes 1 Wings greyish brown without any markings ...... C. griseus Yang & Yang — Wings hyaline or subhyaline, with dark markings ...... 2 2 Hindwings with remarkable blackish-brown markings ...... 3 — Hindwings without dark markings ...... 4 3 Male tenth tergum with apex twisted inward; a sclerite presented beneath male anus ...... C. abnormis Yang & Yang — Male tenth tergum with apex not twisted; no sclerite beneath male anus ...... C. yangi sp. nov. 4 Species from Hainan Island; wing with distinct brown pterostigma ...... C. punctulatus Yang & Yang — Species from Asian mainland; wing without distinct pterostigma ...... 5 5 Male tenth sternum in lateral view with apex not curved dorsad ...... 6 — Male tenth sternum in lateral view with apex curved dorsad ...... 8 6 Forewing with several brownish cloudy markings; male tenth sternum without spine on left lobe ...... 7 — Forewing with four blackish-brown small spots between Rs and M; male tenth sternum with one spine on subapical portion of left lobe ...... C. digitiformis sp. nov. 7 Forewing basally with dark markings; male tenth sternum with apex not curved ventrad in lateral view and somewhat widened in ventral view ...... C. sagittiformis sp. nov. — Forewing basally almost hyaline; male tenth sternum with apex curved ventrad in lateral view and not widened in ventral view ...... C. friedrichi Navás 8 Forewing densely with distinct brownish small spots ...... 9 — Forewing with cloudy brownish markings ...... 10 9 Male tenth sternum with left lobe as long as right lobe, and both lobes curved dorsad in lateral view ...... C. similis sp. nov. — Male tenth sternum with right lobe longer than left lobe, but not curved dorsad in lateral view .....C. nigrovenosus (Weele) 10 Male tenth sternum with left lobe as long as right lobe ...... C. fujianensis Yang & Yang — Male tenth sternum with right lobe longer than left lobe ...... 11 11 Male tenth sternum with right lobe at least twice longer than left lobe ...... C. meridionalis Yang & Yang — Male tenth sternum with right lobe shorter than twice of left lobe ...... C. elongatus sp. nov.

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Fig. 5 A–E. Ctenochauliodes abnormis Yang & Yang. —A. Male genitalia, lateral view. —B. Male tenth tergum, dorsal view. —C. Male tenth tergum, ventral view. —D. Male tenth sternum, ventral view. —E. Sclerite beneath male anus, ventral view. Scale bar = 1 mm.

Ctenchauliodes abnormis Yang & Yang, 1986 (Fig. 5; M8788) strongly sclerotized, from subbasal portion widely divided Ctenchauliodes abnormis Yang & Yang, 1986: 92. into pair of lobes, each with one small spine at tip; in lateral view broad with tip hook-like and curved dorsad; in ventral Material examined. Holotype, CHINA: Guangxi, Longzhou, view basal margin with wide trapezoidal incision, lateral arm Daqingshan, 22°21′N, 106°51′E, 15.v.1963, C. Yang (CAU). triangular, left lobe as long as but slightly wider than right lobe, and distinctly curved outwards. Sclerite beneath anus present, Diagnosis. Wings subhyaline, with distinct blackish-brown semicircular (Fig. 5E), basally with pair of tubercle-like markings. Tenth tergum apically strongly twisted inwards lateral lobes. with ﬂattened tip. Male tenth sternum widely divided into Female: Unknown. pair of lobes from subbasal portion, each with one small spine at tip; left lobe as long as right lobe and distinctly curved Distribution. China (Guangxi). outwards. Remarks. In appearance, this species seems to be closely Description. Male: Body length about 15 mm; forewing related to C. yangi sp. nov. in having similar dark wing length about 24 mm, hindwing length about 21 mm. patterns, but can be easily separated from C. yangi by the Head entirely blackish-brown. Compound eyes brown, male tenth sternum with the left lobe distinctly curved ocelli pale yellow with blackish-brown inner margin. Antenna outwards. In C. yangi, the left lobe is straight and not curved blackish-brown. Mouthparts blackish-brown. outwards. Thorax blackish-brown. Legs brown, with short, dense, yellowish-brown setae. Wings subhyaline, with distinct pale Ctenochauliodes digitiformis sp. nov. (Fig. 6; M8789) blackish-brown markings. Forewings with costal area hyaline Holotype. 1, CHINA: Yunnan, Gengma, Sipaishan, 23°32′N, on basal half and blackish-brown at apex, with tiny blackish- 99°23′E, 1500 m, 3.vi.1980, Huaikuan Qiu (CAU). brown stripes on costal crossveins of basal half; dark markings mostly dotted near crossveins, and often transversely Etymology. The speciﬁc epithet digitiformis refers to the connected, forming narrow bands; apex entirely blackish- elongate and digitiform lateral lobe near the base of the male brown. Hindwings similarly marked, except for bases almost tenth sternum. hyaline and costal area entirely pale black. Veins blackish

brown. Rs four-branched, R2 apically three-branched; Diagnosis. Forewings with pale brown pterostigma and four three crossveins between R1 and Rs; M two-branched; 1 A small blackish-brown spots between Rs and M, and with two-branched. several small brownish spots between branches of Rs. Tenth Abdomen brown. Tenth tergum (Fig. 5A–C) thick at base, tergum with apex strongly twisted ventrad. A pair of slender apically narrowed and strongly twisted inwards, with tip digitiform lateral lobes present near base of male tenth ﬂattened and directed anteriorly. Tenth sternum (Fig. 5A,D) sternum.

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

Fig. 6 A–C. Ctenochauliodes digitiformis sp. nov. —A. Male genitalia, lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth sternum, ventral view. Scale bar = 1 mm.

Description. Male: Body length about 18 mm; forewing length about 24 mm, hindwing length about 21 mm. Head entirely yellowish-brown. Compound eyes blackish- Fig. 7 A–D. Ctenochauliodes elongatus sp. nov. —A. Male genitalia, brown, ocelli yellow with black inner margin. Antenna lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth brown. Mouthparts brown, but labrum and apical portions of sternum, ventral view. —D. Female genitalia, lateral view. Scale maxillary and labial palpi black. bars = 1 mm. Thorax pale brown. Legs brown, with short, dense yellowish setae; apical half of tibiae and tarsi blackish-brown. Wings nearly hyaline with pale brown tips. Forewings with pale and the elongated digitiform lateral lobe near the base of the brown pterostigma and four small blackish-brown spots male tenth sternum. between Rs and M, and with several small brownish spots between branches of Rs. Hindwings without distinct pter- Ctenochauliodes elongatus sp. nov. (Fig. 7; M8753) ostigma, but with two small blackish-brown spots between Rs Holotype. 1, CHINA: Hunan, Nanyue, 27°14′N, 112°43′E, and M. Veins pale yellow except for costal crossveins of 21.vi.1963, Chikun Yang (CAU).

forewings blackish-brown. Rs four-branched, R2 apically three or four-branched; three crossveins between R1 and Rs; Paratypes. 2 2 , same data as holotype (CAU); 1 , CHINA: M two-branched; 1A two-branched. Hunan, Nanyue, Mojingtai, 20.vi.1963, Chikun Yang (CAU). Abdomen blackish brown. Tenth tergum (Fig. 6A,B) short, with apex slightly narrowed and strongly twisted ventrad. Tenth Etymology. The speciﬁc epithet elongatus refers to the elon- sternum (Fig. 6A,C) strongly sclerotized, from subbasal portion gated male tenth tergum and the male tenth sternum. narrowly divided into pair of lobes; in lateral view its apical half subquadrate with spine-like tip; in ventral view basal margin Diagnosis. Forewings greyish brown, usually with cloudy with wide trapezoidal incision, lateral arm subtriangular; left markings transversely connected, forming narrow transverse lobe as long as right lobe and blunt at tip with subapical spine, bands near apex. Male tenth tergum and sternum somewhat right lobe with apical spine. Pair of slender digitiform lateral elongated. Male tenth sternum with left lobe much shorter lobes (Fig. 6A) present near base of male tenth sternum. than right lobe. Female: Unknown. Description. Male: Body length 17–18 mm; forewing length Distribution. China (Yunnan). 22–28 mm, hindwing length 20–22 mm. Head brown with vertex pale brown. Compound eyes Remarks. The new species is distinguished from other brown, ocelli yellow with black inner margin. Antenna blackish Ctenochauliodes species by the wing patterns with four distinct brown. Mouthparts blackish brown, mandibles with apical small blackish-brown spots between Rs and M on the forewings half reddish brown.

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Thorax dark brown. Legs brown, with dense, pale yellow short setae. Wings greyish brown, with many brownish cloudy markings near crossveins, and with pterostgma and apex pale brown. Forewings usually with cloudy markings transversely connected, forming narrow transverse bands near apex. Hindwings paler than forewings, with only two small brownish spots between Rs and M. Veins blackish-

brown. Rs four-branched, R2 apically two-branched; three crossveins between R1 and Rs; M two-branched; 1 A two- branched. Abdomen brown. Tenth tergum (Fig. 7A–B) claviform with apex slightly narrowed; ventral process subtriangular, produced subapically and slightly incurved. Tenth sternum (Fig. 7A,C) strongly sclerotized, from subbasal portion narrowly divided into pair of blade-like lobes; in lateral view its tip long, spine-like and slightly curved dorsad; in ventral view basal margin with shallow trapezoidal incision, lateral arm short and rounded, left lobe somewhat shorter than right lobe and both tips curved inward. Female: Body length 19–20 mm; forewing length about 25–27 mm, hindwing length 22–24 mm. Tenth tergum (Fig. 7D) short and subuliform with blunt tip. Gonocoxite (Fig. 7D) subtriangular, obliquely directed ventrad with acutely pointed tip.

Distribution. China (Hunan). Fig. 8 A–E. Ctenochauliodes friedrichi Navás. —A. Male genitalia, lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth Remarks. See remarks under C. meridionalis. sternum, ventral view. —D. Sclerite beneath male anus, ventral view. —E. Female genitalia, lateral view. Scale bars = 1 mm.

Ctenochauliodes friedrichi Navás, 1932 (Fig. 8; M8754) Ctenochauliodes friedrichi Navás, 1932: 151. Ctenochauliodes forcipatus Kimmins, 1954: 429. syn. nov. 103°19′E, 550–750 m, 22.vi.1957, K. Huang (IZCAS); 2, Emeishan, Qingyinge, 29°32′N, 103°19′E, 800–1000 m, Material examined. 1, CHINA: Sichuan, Kwanshien, 31°00′N, 18.vi.1957, Z. Wang (IZCAS); 1, Jinfoshan, 29°00′N, 107°10′E, 103°36′E, vii.1930, G. Franck (holotype of C. forcipatus) 19.vi.1990, C. Yang (CAU); 1, Guizhou, Zhongpo, 900 m, (NHM); 1, Emeishan, Zhongfengsi, 29°32′N, 103°19′E, 8.vi.1979, Q. Li (CAU); 1, Leishan, Leigongshan, 26°28′N, 16.vi.1990, C. Yang (CAU); 21, Emeishan, 29°32′N, 108°17′E, 1300 m, 29.vi.1988 (IZCAS); 1, Guangxi, Jinxiu, 103°19′E, 29.v.1962 (CAU); 1, Fengdu, Shiping, 29°53′N, Jinzhonggonglu, 24°08′N, 110°11′E, 1100 m, 10.v.1999, H. 107°42′E, 610 m, 2.vi.1994, Y. Zhang (IZCAS); 1, Emeishan, Xiao (IZCAS); 1, Hubei, Xingshan, Longmenhe, 31°13′N, Wanniansi, 29°32′N, 103°19′E, 29.v.1962, Z. Chen (IZCAS); 110°44′E, 1350 m, 18.vii.1993, S. Song (IZCAS); 1, Shen- 11, Emeishan, 29°32′N, 103°19′E, 1800–1900 m, 27/ nongjia, Songbai, 31°45′N, 110°40′E, 23.vi.1977, S. Liu 28.vii1957, Y. Lu (IZCAS); 31, Emeishan, 29°32′N, (CAU); 1, Yunnan, Weixin, 27°51′N, 105°02′E, 1050 m, 103°19′E, 800–1000 m, 11/29.vi.1957, Y. Lu (IZCAS); 1, 12.vi.1980, Z. Tao (CAU); 1, Xishuangbanna, Xiao- Emeishan, Baoguosi, 29°32′N, 103°19′E, 550–750 m, mengyang, 22°01′N, 100°48′E, 850 m, 21.vi.1957, S. Wang 24.v.1957, Z. Wang (IZCAS); 51, Emeishan, Baoguosi, (IZCAS). 29°32′N, 103°19′E, 550–750 m, vi.1957, F. Zhu (IZCAS); 1, Emeishan, 29°32′N, 103°19′E, 600–1200 m, 21.vi.1955 Diagnosis. Forewings hyaline, usually with cloudy markings (IZCAS); 11, Emeishan, 29°32′N, 103°19′E, 580–1150 m, transversely connected, forming narrow transverse bands 21.vi.1955, B. Ou (IZCAS); 1, Emeishan, 29°32′N, 103°19′E, near apex. Male tenth sternum with apical half broad and 900 m, 21.vi.1955-, R. Zhang (IZCAS); 2, Emeishan, slightly curved ventrad, and with rounded tip in lateral view. Qingyinge, 29°32′N, 103°19′E, 800–1000 m, 18.vi.1957, Z. Sclerite beneath male anus present, subtrapezoidal, basally Wang (IZCAS); 12, Emeishan, Baoguosi, 29°32′N, with pair of tubercle-like lateral lobes.

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

Description. Male: Body length 18–23 mm; forewing length 22–30 mm, hindwing length 21–26 smm. Head yellowish brown with blackish-brown clypeus. Compound eyes brown, ocelli yellow with black inner margin. Antenna blackish-brown. Mouthparts blackish-brown. Thorax dark brown. Legs brown, with short, dense pale yellow setae; apices of tibiae and tarsi blackish-brown. Wings hyaline with many brownish cloudy markings near crossveins, and with pterostigma and apex pale brown; sometimes cloudy markings darkened and distinct. Forewings usually with cloudy markings transversely connected, forming narrow transverse bands near apex. Hindwings paler than forewings, mostly without distinct markings, but costal area entirely pale brown, and sometimes several dispersed brownish Fig. 9 A–C. Ctenochauliodes fujianensis Yang & Yang. —A. Male genitalia, markings present along branches of Rs. Rs four-branched, R2 lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth apically three-branched; three crossveins between R1 and Rs; sternum, ventral view. Scale bar = 1 mm. M two-branched; 1A two-branched. Abdomen brown. Tenth tergum (Fig. 8A,B) claviform with apex distinctly narrowed and strongly twisted inward, with Based on a large series in our collections from Sichuan tip directed anteriorly. Tenth sternum (Fig. 8A,C) strongly Province, we found there is only one species, identiﬁed as sclerotized, from subbasal portion narrowly divided into pair C. friedrichi. The features of the male genitalia ﬁt well with of lobes; in lateral view apical half broad with rounded tip, the original illustrations of C. forcipatus. Hence, C. forcipatus slightly curved ventrad; in ventral view basal margin with is treated as a junior synonym of C. friedrichi. trapezoidal incision, lateral arm subtriangular, left lobe suboblong and equal in length to right lobe, right lobe Ctenochauliodes fujianensis Yang & Yang, 1999 (Fig. 9; subtriangular. Sclerite beneath anus (Fig. 8D) present, sub- M8755) trapezoidal, with slightly incised tip, and basally with pair of Ctenochauliodes fujianensis Yang & Yang, 1999: 174. tubercle-like lateral lobes. Female: Body length 25–27 mm; forewing length 30– Material examined. Holotype, CHINA: Fujian, Wuyishan, 35 mm, hindwing length 27–31 mm. Sangang, 27°45′N, 117°40′E, 8.vii.1986, J. Wang & Y. Jiang Tenth tergum (Fig. 8E) short with apical half strongly nar- (CAU). rowed and digitiform. Gonocoxite (Fig. 8E) subtriangular, directed posteriorly. Diagnosis. Forewings with three narrow greyish transverse bands on apical half. Tenth tergum with ventral process Distribution. China (Sichuan, Yunnan, Guizhou, Guangxi, produced from middle portion and slightly curved inwards. Hubei). Male tenth sternum from subbasal portion narrowly divided into pair of acutely pointed lobes, left lobe as long as right Remarks. This species is the most widely distributed species lobe and slightly curved inwards, right lobe straight. of Ctenochauliodes. It can be easily distinguished from the other species by the male tenth sternum with the apex Description. Male: Body length about 12 mm; forewing rounded and curved ventrad, and by subtrapezoidal sclerite length about 19 mm, hindwing length about 17 mm. beneath the male anus. Head blackish brown, with clypeus, postocular margin, The type locality of C. friedrichi is ‘Cinfu Shan’, which and occiput yellowish brown. Compound eyes brown, ocelli must correspond to Mt. Jinfoshan in Sichuan Province. yellow with black inner margin. Antenna blackish brown. Banks (1940) recorded this species from Mt. Emeishan in Mouthparts black. Sichuan Province. In this locality there is another Cteno- Thorax greyish brown, pronotum with anterior margin chauliodes species, C. forcipatus, recorded and described by black. Legs dark brown, with short, dense yellowish setae. Kimmins (1954). In his original description the locality of Wings nearly hyaline with greyish apices and pale grey paratype was cited as ‘Chin-Fu-San’, which should also markings. Forewings with three narrow transverse bands on correspond to Mt. Jinfoshan. According to the original apical half. Hindwings without distinct markings. Veins brown.

illustration of C. friedrichi (only a line ﬁgure of the forewing), Rs four-branched, R2 apically two-branched; three crossveins the wing pattern is remarkably similar to that of C. forcipatus. between R1 and Rs; M two-branched; 1A two-branched.

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Ctenochauliodes moganshanus Yang & Yang, 1992b: 415. syn. nov.

Material examined. Holotype, CHINA: Anhui, Huangshan, Wenquan, 30°08′N, 118°10′E, 24.vii.1977, F. Li (CAU). Paratypes: 1, Anhui, Huangshan, Wenquan, 30°08′N, 118°10′E, 25.vii.1977, F. Li (CAU). 1, Zhejiang, Moganshan, 30°37′N, 119°51′E, 18.vi.1990, J. Lu (CAU); 1, Moganshan, 30°37′N, 119°51′E, 18.vi.1990, Y. Wang (CAU); 1, Moganshan, 30°37′N, 119°51′E, 19.vi.1990, Y. Qiu (CAU).

Diagnosis. Wings greyish brown lacking markings, but median costal areas nearly hyaline in forewings. Male tenth sternum broad with hook-like tip and curved dorsad in lateral view; right lobe longer than left lobe, elongated toward tip, and blade-like in ventral view.

Description. Male: Body length about 14 mm; forewing length about 25 mm, hindwing length about 21 mm. Head brown, with vertex orange. Compound eyes dark brown, ocelli pale yellow with blackish-brown inner margin. Fig. 10 A–C. Ctenochauliodes griseus Yang & Yang. —A. Male genitalia, Antenna black. Mouthparts blackish-brown. lateral view. —B. Male tenth sternum, ventral view. —C. Female genitalia, lateral view. Scale bars = 1 mm. Thorax brown, pronotum with anterior margin black. Legs blackish brown, with short, dense yellowish setae. Wings greyish-brown lacking markings, median costal areas Abdomen pale brown. Tenth tergum (Fig. 9A,B) short and nearly hyaline in forewings. Veins dark brown. Rs four-

subuliform, with ventral digitiform process produced from branched, R2 apically two-branched; three crossveins between middle portion and slightly curved inwards. Tenth sternum R1 and Rs; M two-branched; 1A two-branched. (Fig. 9A,C) strongly sclerotized, from subbasal portion Abdomen dark brown. Tenth tergum (Fig. 10A) broad narrowly separated into pair of blade-like lobes; in lateral with apical half strongly narrowed, ventral process digiti- view broad with apical half distinctly pointed and curved form, produced from subapical portion and curved inwards. dorsad; in ventral view basal margin with wide trapezoidal Tenth sternum (Fig. 10A,B) strongly sclerotized, from incision, lateral arm nearly rounded, left lobe as long as right subbasal portion narrowly divided into pair of blade-like lobe and slightly curved inwards with apical 1/3 pointed, lobes; in lateral view broad with tip hook-like and curved right lobe straight. dorsad; in ventral view basal margin with wide trapezoidal Female: Unknown. incision, lateral arm nearly rounded, left lobe incurved, right lobe longer than left lobe. Distribution. China (Fujian). Female: Body length 17–23 mm; forewing length 27– 31 mm, hindwing length 23–26 mm. Remarks. This species appears to be closely related to Tenth tergum (Fig. 10C) with straight dorsal margin, apex C. griseus in having the similar ventral process of the male narrowed and digitiform. Gonocoxite (Fig. 10C) broad and tenth tergum and the male tenth sternum not narrowed subtriangular. behind lateral arm in lateral view, but can be easily separated from C. griseus by the wing pattern with sparse greyish mark- Distribution. China (Anhui, Zhejiang). ings, and by the male tenth sternum with the left lobe as long as the right lobe. In C. griseus, the wing is greyish without any Remarks. This species is distinguished from other species of markings, and the left lobe of the male tenth sternum is Ctenochauliodes by the greyish wings lacking markings. After shorter than the right lobe. careful examination of the types of C. moganshanus, the general appearance as well as the female genitalia ﬁt well with Ctenchauliodes griseus Yang & Yang, 1992 (Fig. 10; M8756) that of C. griseus. No signiﬁcant differences are found to Ctenchauliodes griseus Yang & Yang, 1992a: 1. separate these two species at present. Hence, C. moganshanus

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

Head blackish-brown with vertex pale yellow. Compound eyes dark brown, ocelli yellow with blackish inner margin. Antenna blackish-brown. Mouthparts blackish-brown. Thorax blackish-brown, with meso- and metanotum greyish- brown. Legs brown, with short, dense yellowish setae. Wings hyaline with many cloudy brownish markings near crossveins, and with pterostigma and apex pale brown. Forewings usually with cloudy markings transversely connected, forming narrow transverse bands near apex. Hindwings paler than forewings, lacking distinct brownish markings. Veins

blackish-brown. Rs four-branched, R2 apically two-branched; three crossveins between R1 and Rs; M two-branched; 1A two-branched. Abdomen blackish-brown. Tenth tergum (Fig. 11A,B) short and subuliform; in lateral view apex distinctly narrowed, ventral process subtriangular and curved inwards. Tenth sternum (Fig. 11A,C) strongly sclerotized, from subbasal portion narrowly divided into pair of lobes; in ventral view basal margin with wide trapezoidal incision, lateral arm nearly rounded; left lobe short and broad with spine-like dorsal apical corner in lateral view, apical half slightly incurved in ventral view; right lobe at least twice as long as left lobe, in lateral view apical half strongly narrowed, long spine-like, and obliquely produced dorsad, in ventral view Fig. 11 A–E. Ctenochauliodes meridionalis Yang & Yang. —A. Male blade-like with tip slightly incurved. Sclerite beneath anus genitalia, lateral view. —B. Male tenth tergum, ventral view. —C. Male present (Fig. 11D), in ventral view trilobed with truncate tenth sternum, ventral view. —D. Sclerite beneath male anus, ventral apical margin. view. —E. Female genitalia, lateral view. Scale bars = 1 mm. Female: Body length 17–18 mm; forewing length 26– 29 mm, hindwing length about 23–26 mm. Tenth tergum (Fig. 11E) with arched dorsal margin, apical is considered to be conspeciﬁc with C. griseus and herein half distinctly narrowed and slightly curved ventrad. Gono- designated as a junior synonym of C. griseus. coxite (Fig. 11E) subtriangular, obliquely directed dorsad, with acutely pointed tip. Ctenchauliodes meridionalis Yang & Yang, 1986 (Fig. 11; M8757) Distribution. China (Guangxi). Ctenchauliodes meridionalis Yang & Yang, 1986: 93. Remarks. This species appears to be closely related to Material examined. Holotype, CHINA: Guangxi, Wuming, C. elongatus in its similar wing pattern, but can be easily Damingshan, 23°27′N, 108°30′E, 21.v.1963, C. Yang (CAU). distinguished from the latter by the male tenth sternum with Allotype, same data as holotype (CAU). Paratypes: 13, the right lobe more than twice as long as the left lobe. In Guangxi, Wuming, Damingshan, 23°27′N, 108°30′E, 22/ C. elongatus, both lobes of the male tenth sternum are elongated 23.v.1963 (CAU); 1, Longsheng, Tianpingshan, 25°28′N, and its right lobe is less than twice as long as the left lobe. 109°57′E, 8.vi.1963, C. Yang (CAU). Ctenochauliodes nigrovenosus (van der Weele, 1907) (Fig. 12; Diagnosis. Forewings usually with cloudy markings trans- M8758) versely connected, forming narrow transverse bands near Chauliodes nigrovenosus van der Weele, 1907: 257. apex. Male tenth sternum with right lobe at least twice as long as left lobe. Sclerite beneath male anus present, in ventral Material examined. 1, CHINA: Guangxi, Huaping, 25°28′N, view trilobed with truncate apical margin. 109°57′E, 24.vi.1982, C. Yang (CAU); 11, Huaping, Tianpingshan, 25°28′N, 109°57′E, 5.vi.1963, C. Yang (CAU); Description. Male: Body length 16–18 mm; forewing length 1, Wuming, Damingshan, 23°27′N, 108°30′E, 21.v.1963, 25–28 mm, hindwing length 22–24 mm. C. Yang (CAU).

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Fig. 13 Ctenochauliodes punctulatus Yang & Yang — Female genitalia, lateral view. Scale bar = 1 mm.

of lobes; in ventral view basal margin with shallow trapezoidal incision, lateral arm nearly rounded; left lobe slightly shorter Fig. 12 A–E. Ctenochauliodes nigrovenosus (van der Weele) —A. Male than right lobe, in lateral view with tip hook-like and curved genitalia, lateral view. —B. Male tenth tergum, ventral view. —C. Male dorsad, in ventral view with tip incurved; right lobe in lateral tenth sternum, ventral view. —D. Sclerite beneath male anus, ventral view. —E. Female genitalia, lateral view. Scale bars = 1 mm. view with tip spine-like but not curved dorsad, in ventral view blade-like with tip incurved. Sclerite beneath anus present (Fig. 12D), trilobed, with wide median lobe triangular, with Diagnosis. Forewings densely dotted, with several narrow lateral lobes slender and acutely pointed. transverse bands on apical half. Male tenth sternum in lateral Female: Body length 25–27 mm; forewing length 30– view with apex of left lobe curved dorsad, right lobe much 35 mm, hindwing length 27–31 mm. longer but not curved dorsad. Tenth tergum (Fig. 12E) short and subuliform with blunt tip. Gonocoxite (Fig. 12E) subtriangular, obliquely directed dorsad. Description. Male: Body length 18–21 mm; forewing length 26–27 mm, hindwing length 23–24 mm. Distribution. China (Guangxi); Vietnam. Head yellowish-brown with frons pale brown. Compound eyes brown, ocelli yellow with blackish-brown inner margin. Remarks. This species was established by van der Weele Antenna blackish-brown. Mouthparts blackish-brown. (1907) based on two female specimens from ‘Mauson Moun- Thorax pale brown. Legs yellowish-brown, with short, tains’ in Vietnam. According to the original descriptions and dense, yellowish setae; apices of tibiae and tarsi pale brown. the habitus photo, we identiﬁed a series of specimens from Wings hyaline, with many brownish markings, and with ptero- Guangxi Province, which is adjacent to Vietnam, as stigma and apex extremely pale brown. Forewings densely C. nigrovenosus. Consequently, the male is ﬁrstly described dotted, with several narrow transverse bands on apical half. herein. This species appears closely related to C. similis in Hindwings nearly hyaline except for costal area mostly having similar wing patterns of dense spots on the forewings, pale brown, and apex with some indistinct small spots. Rs four- but can be easily separated from C. similis by the male tenth

branched, R2 apically two or three-branched; three crossveins sternum with the right lobe not curved dorsad in lateral view. between R1 and Rs; M two-branched; 1A two-branched. In C. similis, the right lobe of the male tenth sternum is Abdomen brown. Tenth tergum (Fig. 12A,B) short clavi- curved dorsad in lateral view. form with apical half distinctly narrowed and curved ventrad; ventral process subtriangular, produced from subapical portion Ctenchauliodes punctulatus Yang & Yang, 1990 (Fig. 13; and slightly incurved. Tenth sternum (Fig. 12A,C) strongly M8759) sclerotized, from subbasal portion narrowly separated into pair Ctenchauliodes punctulatus Yang & Yang, 1990: 99.

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

Material examined. Holotype, CHINA: Hainan, Jianfengling, 18°42′N, 108°47′E, 7.iv.1982, Y. Liu (CAU). Paratype 1, same data as holotype (CAU). 1, 1000 m, 23.iv.1980, S. Wang (IZCAS).

Diagnosis. Forewings with distinct brown pterostigma, and with many small brownish spots densely dispersed along longitudinal veins. Hindwings without any brownish spots except for short brownish pterostigma.

Description. Female: Body length 19–21 mm; forewing length 24–26 mm, hindwing length 20–23 mm. Head greyish-brown with vertex yellowish brown. Com- pound eyes brown, ocelli yellow with brown inner margin. Fig. 14 A–C. Ctenochauliodes sagittiformis sp. nov. —A. Male genitalia, Antenna blackish-brown. Mouthparts pale brown, but lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth mandibles yellow with apical half pale reddish-brown. sternum, ventral view. Scale bar = 1 mm. Thorax pale brown, meso- and metanotum each with centre yellowish-brown. Legs yellow, with short, dense yellow setae; tips of tibiae and tarsi brown. Wings hyaline. Forewings arrow-like in lateral view, middle portion bluntly produced with distinct brown pterostigma; many small brownish spots dorsad and acutely produced ventrad. densely dispersed along longitudinal veins. Hindwings with pterostigma much shorter and without any brownish spots. Description. Male: Body length about 14 mm; forewing length

Veins yellowish-brown. Rs four-branched, R2 apically three about 22 mm, hindwing length about 21 mm. or four-branched; three crossveins between R1 and Rs; M Head blackish brown with vertex pale brown. Compound two-branched; 1A two-branched. eyes brown, ocelli yellow with black inner margin. Antenna Abdomen yellowish brown. Tenth tergum (Fig. 13) short blackish-brown. Mouthparts brown. and subuliform with blunt tip. Gonocoxite (Fig. 13) broad Thorax blackish-brown. Legs yellowish-brown, with with arched ventral margin and narrowed tip. short, dense, yellowish setae; apical half of tibiae and tarsi Male: Unknown. dark brown, tarsal claws pale red. Wings pale smoky brown with brownish cloudy markings. Forewings with costal area Distribution. China (Hainan). pale brown except for four or ﬁve hyaline costal cellules on middle portion; cloudy markings dotted throughout and Remarks. This species was established based on two female much darker on basal half. Hindwings mostly without any specimens from Hainan, which is the second largest island of markings, but with brownish spots on costal cellules. Rs

China. Though the male is still unknown, this species is four-branched, R2 apically three-branched; three crossveins distinguished from the other species of Ctenochauliodes by the between R1 and Rs; M two-branched; 1A two-branched. wing pattern with dense brownish spots and distinct brown Abdomen yellowish-brown, dorsally with black vittae near pterostigma. lateral sides. Tenth tergum (Fig. 14A,B) moderately elongated with apical half slightly narrowed and curved ventrad, Ctenochauliodes sagittiformis sp. nov. (Fig. 14; M8760) tip twisted inwards. Tenth sternum (Fig. 14A,C) strongly Holotype. 1, CHINA: Guangdong, Ruyuan, Nanling, sclerotized, from subbasal portion narrowly divided into pair 23°20′N, 115°23′E, 9/18.vii.2005 (CAU). of lobes; in lateral view arrow-like, middle portion bluntly produced dorsad and acutely produced ventrad; in ventral Paratype. 1, CHINA: Fujian, Chong-an, Xingcun, San- view basal margin with wide trapezoidal incision, lateral arm gang, 27°45′N, 117°40′E, 740 m, 26.vii.1960, Yiran Zhang elongate and subtriangular; left lobe as long as but much wider (IZCAS). than right lobe, both lobes with apical half slightly widened. Female: Unknown. Etymology. The speciﬁc epithet sagittiformis refers to the arrow-like male tenth sternum in lateral view. Distribution. China (Guangdong, Fujian).

Diagnosis. Forewings with cloudy markings dotted through- Remarks. The new species is distinguished from other species out and much darker on basal half. Male tenth sternum of Ctenochauliodes by the wing pattern where the forewings are

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

sternum with left lobe as long as right lobe, and both lobes broad with apical half distinctly narrowed and incurved.

Description. Male: Body length 18–23 mm; forewing length 24–28 mm, hindwing length 21–25 mm. Head brown with clypeus and middle portion of vertex yellowish-brown. Compound eyes brown, ocelli yellow with black inner margin. Antenna blackish-brown. Mouthparts blackish-brown. Thorax brown. Legs yellowish-brown, with short, dense, yellowish setae; apices of tibiae and tarsi pale brown. Wings hyaline, with many brownish marks, and with pterostgma extremely pale brown. Forewings densely dotted throughout except for one transverse hyaline area on middle portion. Hindwings nearly hyaline except for costal area mostly pale brown, apex densely dotted with small brownish spots.

Rs four-branched, R2 apically two- or three-branched; three crossveins between R1 and Rs; M two-branched; 1A two-branched. Abdomen brown. Tenth tergum (Fig. 15A,B) short claviform with apical half distinctly narrowed and curved ventrad; ventral process subtriangular, produced from subapical portion, and distinctly incurved. Tenth sternum (Fig. 15A,C) strongly sclerotized, from subbasal portion narrowly separated into pair of blade-like lobes; in lateral view apical Fig. 15 A–E. Ctenochauliodes similis sp. nov. —A. Male genitalia, half broad with tip spine-like and curved dorsad; in ventral lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth sternum, ventral view. —D. Sclerite beneath male anus, ventral view. view basal margin with a trapezoidal incision, lateral arm —E. Female genitalia, lateral view. Scale bars = 1 mm. nearly rounded; left lobe as long as right lobe, and both lobes broad with apical half distinctly narrowed and incurved. Sclerite beneath anus present (Fig. 15D), trilobed, much darker on the basal portions. The features of the male with rounded median lobe and acutely pointed lateral genitalia are also peculiar with its moderately elongated tenth lobes. tergum with the tip somewhat twisted inwards, and by the Female: Body length 20–22 mm; forewing length 24– arrow-like male tenth sternum with the blunt tip. 31 mm, hindwing length 21–27 mm. Tenth tergum (Fig. 15E) short, digitiform, and slightly Ctenochauliodes similis sp. nov. (Fig. 15; M8761) curved ventrad. Gonocoxite (Fig. 15E) subtriangular, obliquely Holotype. 1, CHINA: Guangxi, Wuming, Damingshan, directed dorsad; left lobe with tip much blunter than right 21.v.1963, Chikun Yang (CAU). lobe.

Paratypes. 5, same data as holotype (CAU); 25, Guangxi, Distribution. China (Guangxi). Wuming, Damingshan, 23°27′N, 108°30′E, 22.v.1963, Chikun Yang (CAU); 1, Wuming, Damingshan, 23°27′N, Remarks. See the remarks on C. nigrovenosus. 108°30′E, 23.v.1963, Chikun Yang (CAU); 1, Huaping, Tianpingshan, 25°28′N, 109°57′E, 6.vi.1963, Chikun Yang Ctenochauliodes yangi sp. nov. (Fig. 16; M8762) (CAU); 1, Huaping, 25°28′N, 109°57′E, 25.vi.1982, Chikun Holotype. 1, CHINA: Guangxi, Fangcheng, Fulong, Yang (CAU). 21°49′N, 107°57′E, 500 m, 24.v.1999, Dajun Liu (IZCAS).

Etymology. The speciﬁc epithet similis refers to its similar Paratypes. 1, same data as holotype (IZCAS); 1, CHINA: appearance to C. nigrovenosus. Guangxi, Fangcheng, Banbaxiang, 21°49′N, 107°57′E, 550 m, 4.vi.2000, Jian Yao (IZCAS); 1, CHINA: Guangdong, Diagnosis. Forewing densely dotted throughout except for Dinghushan, 23°11′N, 112°31′E, 18.vi.1983, Peiyu Wu one transverse hyaline area on middle portion. Male tenth (IZCAS).

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

brown stripes; most marks dotted near crossveins and transversely connected, forming several wide transverse bands; basal transverse bands short, median transverse band V-shaped, apical transverse bands somewhat shortened and narrowed. Hindwings similarly marked, with basal transverse bands much wider, and with two apical transverse bands

much narrower and nearly parallel. Rs four-branched, R2 apically two-branched; three crossveins between R1 and Rs; M two-branched; 1A two-branched. Abdomen blackish brown. Tenth tergum (Fig. 16A,B) with straight dorsal margin, apex slightly narrowed with blunt tip; ventral process feebly developed and invisible in lateral view. Tenth sternum (Fig. 16A,C) strongly sclerotized, from subbasal portion widely divided into a pair of apically pointed lobes; in lateral view apical half broad, triangular, slightly curved dorsad, ventrally produced triangularly; in ventral view basal margin with a wide trapezoidal incision, lateral arm subtriangular; left lobe as long as but much thinner than right lobe, and both lobes straightly directed and not incurved. Sclerite beneath anus present (Fig. 16D), trilobed, basal margin with a deep trapezoidal incision, median lobe with apical margin truncate, lateral lobes acutely produced. Female: Body length 23–25 mm; forewing length 28– 30 mm, hindwing length 26–28 mm. Fig. 16 A–F. Ctenochauliodes yangi sp. nov. —A. Male genitalia, Tenth tergum (Fig. 16E) subtriangular with blunt tip. lateral view. —B. Male tenth tergum, ventral view. —C. Male tenth Gonocoxite (Fig. 16E,F) in lateral view subtriangular, with sternum, ventral view. —D. Sclerite beneath male anus, ventral view. —E. Female genitalia, lateral view. —F. Female gonocoxite, ventral pair of small digitiform processes; in ventral view asymmetric, view. Scale bars = 1 mm. left lobe with outer margin vertically incurved at middle portion, right lobe with outer margin much shorter and indistinctly incurved, and only with tip somewhat arcuately curved. Etymology. The speciﬁc epithet yangi refers to the excellent entomologist, Professor Chikun Yang, who passed away in Distribution. China (Guangxi, Guangdong). 2006, in honour of his contribution to the systematics of the Chinese Neuropterida. Remarks. See the remarks on C. abnormis.

Diagnosis. Wings subhyaline with distinct blackish-brown Acknowledgements marks. Male tenth sternum in lateral view with apical half We thank Mr Jian Yao (Chinese Academy of Science, Beijing) broad and triangular; in ventral view left lobe as long as but for providing many specimens from the insect collections. Our much thinner than right lobe, and both lobes straightly thanks also go to Dr Fumio Hayashi (Tokyo Metropolitan directed and not incurved. University, Tokyo) and Dr Oliver S. Flint, Jr. (Washington, D. C) for offering many references and specimens. We are much Description. Male: Body length 19–21 mm; forewing length indebted to Dr Bradley J. Sinclair (Bonn) and two anonymous 25–27 mm, hindwing length 22–25 mm. referees for their attention to the manuscript. This research Head blackish-brown. Compound eyes brown, ocelli yellow was supported by the National Natural Science Foundation with blackish-brown inner margin. Antenna blackish-brown. of China (30370174, 30225009, 30200025, 30430100). Mouthparts blackish-brown, mandibles with apices reddish- brown. References Thorax blackish-brown. Legs blackish-brown, with dense, Banks, N. (1940). Report on certain groups of Neuropteroid insects pale brown, short setae. Wings subhyaline, with distinct from Szechwan, China. Proceedings of the United States National blackish-brown marks, and with apex blackish-brown. Museum, 88 (3079), 173–220. Forewings with costal areas hyaline except for apices entirely Contreras-Ramos, A. (2004). Is the family Corydalidae (Neuropterida, blackish-brown, costal crossveins marked with tiny blackish- Megaloptera) a monophylum? Denisia, 13, 135–140.

X. Y. Liu & D. Yang • Revision of the Ctenochauliodes species

Eriksson, T. (1998). Autodecay, Version 4.0 (program distributed by species. The pectinate female antenna is considered to be the author). Stockholm: Royal Swedish Academy of Sciences, synapomorphic for the genus Ctenochauliodes. Bergius Foundation. Kimmins, D. E. (1954). A new genus and some new species of the 3 Male tenth tergum: (0) not twisted apically; (1) twisted Chauliodini (Megaloptera). Bulletin of the British Museum (Natural History), Entomology, 3, 415–444. apically. In most species of Chauliodinae, the male tenth Liu, X. Y. & Yang, D. (2005). Revision of the Protohermes changningensis tergum is normally directed with the apex not twisted, while species group from China (Megaloptera: Corydalidae: Corydalinae). it is distinctly twisted apically in C. abnormis, C. digitiformis, Aquatic Insects, 27, 167–178. C. friedrichi, and C. sagittiformis (Figs 5B, 6B, 8B and 14B). Liu, X. Y. & Yang, D. (2006a). Revision of the genus Sialis from Oriental China (Megaloptera: Sialidae). Zootaxa, 1108, 23–35. 4 Male tenth tergum: (0) not curved ventrad; (1) curved Liu, X. Y. & Yang, D. (2006b). Phylogeny of the subfamily of Chaulio- ventrad. The male tenth tergum is normally not curved dinae (Megaloptera: Corydalidae), with description of a new genus from the Oriental Realm. Systematic Entomology, in press. ventrad except for C. digitiformis, where the apex is strongly Liu, X. Y., Yang, D., Ge, S. Q. & Yang, X. K. (2005). Phylogenetic curved ventrad (Fig. 6B). review of the Chinese species of Acanthacorydalis (Megaloptera: Corydalidae). Zoologica Scripta, 34, 373–387. 5 Male tenth tergum in ventral view with tip: (0) not directed Maddison, D. R. & Maddision, W. P. (2000). MacClade, Version 4.0. anteriorly; (1) directed anteriorly. In C. abnomis and C. friedrichi, Sunderland, MA: Sinauer Associates. the tip of the male tenth tergum is anteriorly directed in Navás, L. (1932). Decadas de insectos nuevos. Brotéria, 1, 145–155. ventral view, while in the other Ctenochauliodes species and the Penny, N. D. (1993). The phylogenetic position of Chloroniella peringueyi (Megaloptera: Corydalidae) and its zoogeographic outgroups it is not directed anteriorly (Figs 5B,C and 8B). significance. Entomological News, 104, 17–30. Swofford, D. L. (2002). PAUP* — Phylogenetic Analysis Using Parsimony 6 Male tenth tergum, ventral process: (0) absent; (1) present. (*and Other Methods), Version 4.0b10. Sunderland, MA: Sinauer In the outgroups, C. abnomis, C. digitiformis, C. friedrichi and Associate. C. sagittiformis, the male tenth tergum is simple without any van der Weele, H. W. (1907). Notizen ueber Sialiden und Besch- process at the subapical portion, while in the other Cteno- reibung einiger neuen Arten. Notes from the Leyden Museum, 28, chauliodes species, there is a process ventrally produced from 227–264. van der Weele, H. W. (1909). New genera and species of Megaloptera the subapical portion of the male tenth tergum (Figs 7B, 9B, Latr. Notes from the Leyden Museum, 30, 249–264. 10A, 11B, 12B, 15B and 16B). van der Weele, H. W. (1910). Megaloptera Monographic Revision. Collections Zoologiques du Baron Edm. de Selys Longchamps, 5, 1–93. 7 Male tenth tergum, ventral process: (0) feebly developed; (1) Yang, C. K. & Yang, D. (1986). New fishflies from Guangxi, China strongly developed. In C. elongatus, C. fujianensis, C. griseus, (Megaloptera: Corydalidae). Entomotaxonomia, 8, 85–95. C. meridionalis, C. nigrovenosus, and C. similis, the ventral process Yang, C. K. & Yang, D. (1990). The fishflies from Hainan Island of the tenth tergum is strongly developed (Figs 7B, 9B, 10A, (Megaloptera: Corydalidae). Acta Zootaxonomica Sinica, 15, 98–100. Yang, C. K. & Yang, D. (1992a). Two new species of Corydalidae 11B, 12B and 15B), while in C. yangi it is rather feebly (Megaloptera). Entomological Journal of East China, 1, 1–3. developed (Fig. 16B). Yang, C. K. & Yang, D. (1992b). Megaloptera of Mount Mogan with one new species. Journal of Zhejiang Forestry Colleage, 9, 414–417. 8 Male tenth tergum, ventral process: (0) not triangular; (1) Yang, C. K. & Yang, D. (1999). Megaloptera, Corydalidae. In triangular. In C. elongatus, C. meridionalis, C. nigrovenosus and B. K. Huang (Ed.) Fauna of Insects in Fujian Province of China, Vol. 3 C. similis, the ventral process of the tenth tergum is triangular (pp. 168–176). Fuzhou: Fujian Science Press. (Figs 7B, 11B, 12B and 15B), while in C. fujianensis and Zhang, Y. Z. (1998). The second revision of zoogeographical regions of China. Acta Zootaxonomica Sinica, 23 (Suppl.), 207–222. C. griseus it is digitiform (Figs 9B and 10A).

Appendix 1 9 Male tenth tergum with tip: (0) blunt; (1) acutely pointed. Characters and character states employed in the cladistic In C. elongatus, C. meridionalis, C. nigrovenosus and C. similis, analysis of the Ctenochauliodes species. Character states were the tip of the male tenth tergum is acutely pointed in ventral scored 0–2, ? (0 = plesiomorphic state, 1–2 = apomorphic view (Figs 7B, 11B, 12B and 15B), while it has a blunt tip in state, ? = state unknown). the other Ctenochauliodes species and the outgroups.

1 Male antenna: (0) filiform; (1) pectinate. The male antenna is 10 Male tenth tergum with spinous setae: (0) dense; (1) sparse. filiform in the outgroups, pectinate in the Ctenochauliodes species. In most species of Chauliodinae, the male tenth tegum bears dense spinous setae. But in the Ctenochauliodes species, the 2 Female antenna: (0) filiform; (1) pectinate. The female antenna spinous setae are rather stout and sparse, which is considered is filiform in the outgroups, pectinate in the Ctenochauliodes to be a synapomorphic character of the genus Ctenochauliodes.

Revision of the Ctenochauliodes species • X. Y. Liu & D. Yang

11 Male tenth sternum: (0) bifurcated from the middle C. meridionalis, C. nigrovenosus and C. similis, it is blade-like portion; (1) bifurcated from the subbasal portion. The with an acutely pointed tip in ventral view (Figs 7C, 9C, 10C, bifurcated male tenth sternum is shared by several genera of 11C, 12C and 15C). In the other Ctenochauliodes species and Chauliodinae, such as Archichauliodes, Ctenochauliodes, the outgroups, it is blunt in ventral view. Dysmicohermes, Orohermes, and Protochauliodes. The furcation is from the middle portion of the male tenth sternum, 20 Male tenth sternum, lateral arm in ventral view: (0) more or forming a relatively shallow incision, which is considered to less elongated and subtriangular; (1) short and rounded. The be plesiomorphic. In Ctenochauliodes, the male tenth sternum lateral arms of the male tenth sternum are the joints of the is bifurcated from the subbasal portion, forming a rather deep male tenth sternum to connect with the ninth abdominal incision, which is synapomorphic for the genus. segment. In the outgroup species the lateral arm is somewhat elongated and subtriangular. In most Ctenochauliodes species, 12 Male tenth sternum with pair of lobes formed by median the lateral lobe is shortened and rounded, except for incision: (0) symmetric; (1) asymmetric. The furcation of the C. abnormis, C. digitiformis, C. friedrichi, C. sagittiformis and male tenth sternum forms a pair of lobes which are variously C. yangi (Figs 5D, 6C, 8C and 14C). modiﬁed. The lobes are plesiomorphically symmetric in many genera of Chauliodinae, but in Ctenochauliodes they are 21 Male tenth sternum in lateral view: (0) narrowed behind asymmetric in shape and even in length (see char. 18). lateral arm; (1) not narrowed behind lateral arm. In the outgroups and the most Ctenochauliodes species, the male 13 Male tenth sternum: (0) curved dorsad; (1) not curved dorsad. tenth sternum is narrowed behind the lateral arm and then The male tenth sternum in most Chauliodinae is curved dorsad, distinctly widened in lateral view, while in C. fujianensis and while it is directed posteriorly and not curved dorsad in C. C. griseus, the narrowed link of the male tenth sternum is digitiformis, C. friedrichi and C. sagittiformis (Figs 6A, 8A and 14A). absent between the lateral arm and the widened median portion (Figs 9A and 10A). 14 Male tenth sternum, left lobe in ventral view with tip: (0) blunt; (1) acutely pointed. In the outgroups, C. digitiformis, 22 Sclerite between anus and the male tenth sternum: (0) C. friedrichi and C. sagittiformis, the left lobe of the male tenth absent; (1) present but not trilobed; (2) present and trilobed. sternum is blunt at the tip (Figs 6C, 8C and 14C), while in the In some Ctenochauliodes species, there is a sclerotized plate other Ctenochauliodes species it is acutely pointed at the tip. beneath the anus. Kimmins (1954) presumed this plate might be the tenth sternum. Herein, we follow the terminology of 15 Male tenth sternum, left lobe in ventral view: (0) not curved Megaloptera used by Contreras-Ramos (2004), so the outward; (1) curved outward. aedeagus of Chauliodinae (Kimmins 1954) is considered to be the male tenth sternum. Thus, the sclerite between the 16 Male tenth sternum, left lobe in ventral view: (0) not anus and the male tenth sternum might be the remnant of the widened apically; (1) widened apically. paramere, which might be secondarily derived. In C. abnormis and C. friedrichi, this sclerite is nearly semicircular or 17 Male tenth sternum, left lobe in ventral view: (0) without spine subtrapezoidal but not trilobed (Figs 5E and 8D), while in on subapical portion; (1) with a spine on subapical portion. C. meridionalis, C. nigrovenosus, C. similis and C. yangi, it is trilobed (Figs 11D, 12D, 15D and 16D). 18 Male tenth sternum, right lobe: (0) as long as left lobe; (1) much longer than left lobe. In the outgroups, the male tenth 23 Male tenth sternum, lateral lobe: (0) absent; (1) present and sternum is bifurcated into a pair of symmetric lobes, but in digitiform; (2) present and tubercle-like. In C. abnormis, C. elongatus, C. griseus, C. meridionalis and C. nigrovenosusit is C. digitiformis and C. friedrichi, there is a pair of lobes near the asymmetrically bifurcated and the right lobe is longer than base of the male tenth sternum or fused with the sclerite the left process (Figs 7C, 10C, 11C and 12C). In the other beneath the anus, which is considered to be apomorphic and Ctenochauliodes species, the right lobe is as long as the left homogeneous with the lateral lobes of the male tenth lobe, even though it is different from the left lobe in shape. sternum in Corydalinae. In C. abnormis and C. friedrichi, the lateral lobes are fused with the basal lateral corners of the 19 Male tenth sternum, right lobe in ventral view: (0) blunt at sclerite beneath the anus (Figs 5E and 8D), while in tip; (1) with one small spine at tip; (2) with acutely pointed C. digitiformis they are rather long and digitiform (Fig. 6A). tip. In C. abnormis and C. digitiformis, the tip of the right lobe of the male tenth sternum bears a small spine in ventral view 24 Female gonocoxite: (0) bluntly tapered toward tip; (1) (Figs 5D and 6C), while in C. elongatus, C. fujianensis, C. griseus, acutely tapered toward tip; (2) with tip incised ventrally.