Systematics of the Protohermes Xanthodes Species-Group in Eastern Asia (Megaloptera: Corydalidae)

Entomological Science (2006) 9, 399–409 doi:10.1111/j.1479-8298.2006.00186.x

ORIGINAL ARTICLE

Systematics of the Protohermes xanthodes species-group in eastern Asia (Megaloptera: Corydalidae)

Xingyue LIU1, Fumio HAYASHI2 and Ding YANG1 1Department of Entomology, China Agricultural University, Beijing, China; and 2Department of Biology, Tokyo Metropolitan University, Minamiosawa, Hachioji, Tokyo, Japan

Abstract A new species-group of the dobsonﬂy genus Protohermes is proposed, the Protohermes xanthodes species- group. Three species from eastern Asia belonging to the new species-group are redescribed and illustrated. Phylogenetic relationships among the species in this group, as well as the biogeography of these species, are discussed on the basis of a cladistic analysis. Key words: Corydalinae, new species-group, phylogeny, Protohermes, taxonomy.

INTRODUCTION status by Contreras-Ramos (1998). Recent studies of the systematics of Protohermes have been carried out by The Asian dobsonfly genus Protohermes is one of the Hayashi (1989a,b, 1990), Yang (1985), Yang and Yang most species-rich genera within the subfamily Corydali- (1988), and Liu and Yang (2005, 2006a,b). nae. It includes up to 40 described species, which are Previously, two Protohermes species-groups, the predominantly distributed in the Oriental region. Adult P. changninganus species-group and the P. davidi spe- Protohermes are characterized by a yellowish body with cies-group, have been recognized (Liu & Yang 2005, various blackish marks on the head and pronotum, 2006b). The P. costalis species-group was recognized as wings usually with several yellowish or whitish marks, the third species-group by X. Liu, F. Hayashi and D. forewing with a three-branched 1A, and a posteriorly Yang (unpubl. data, 2006). In the present paper, a fourth incised male ninth sternum. The male tenth tergite varies species-group, the Protohermes xanthodes species- generally from an elongate band and flattened valve, to group, is proposed, comprising the species P. xanthodes, a short subcylindrical or claviform tube. Early revisions P. grandis, and P. immaculatus. All three species are of the genus were made by European neuropterologists keyed, redescribed, and reillustrated herein. The first (van der Weele 1910; Lestage 1927; Kimmins 1948). record of the genus Protohermes from Korea is This genus, together with Neurhermes Navás, were included, as well as a discussion of the phylogeny and established as a well-supported monophyletic group biogeography of the P. xanthodes species-group. according to cladistic analyses (Glorioso 1981; Penny 1993; Contreras-Ramos 1998). The group was thought to be a sister group to the remaining genera within the MATERIALS AND METHODS Corydalinae by Penny (1993), or a sister group to the Preparation of specimens remaining Corydalinae genera except Chloroniella Esben-Petersen with the most primitive phylogenetic Specimens were collected mostly using light traps in mountainous regions at sites close to clean running water. Genitalic preparations were made by clearing the apex of the abdomen in a saturated KOH solution for Correspondence: Dr Ding Yang, Department of Entomology, 8–10 h. The apex of the abdomen was transferred to China Agricultural University, Beijing 100094, China. glycerin for further dissection and examination, then Email: [email protected] moved to fresh glycerin and stored in a microvial pinned Received 22 December 2005; accepted 14 August 2006. below the specimen.

The specimens for the present study are deposited at 6 Male tenth tergite: without setal tuft, 0; with setal the Entomological Museum of the China Agricultural tuft, 1. University (CAU), Beijing; the Institute of Zoology, 7 Ninth gonostylus: slender, 0; thick, 1. Chinese Academy of Sciences (IZCAS), Beijing; the 8 Male tenth sternite with median portion: simple, 0; Shanghai Entomological Museum, Chinese Academy of enlarged and somewhat modified, 1. Sciences (SEMCAS), Shanghai; and Fumio Hayashi’s 9 Male tenth sternite with lateral lobes: nearly membra- personal collection (HC), Tokyo, Japan. nous, 0; moderately sclerotized, 1. Morphological terminology generally follows Glori- The present cladistic analysis was performed using oso (1981) except for the terminology for the tenth PAUP* version 4.0b10 (Swofford 2002) by using heuris- abdominal segment, which follows Contreras-Ramos tic parsimony analysis, with 1000 random stepwise (1998). additions of taxa (TBR branch swapping) under ACCT- RAN optimization, characters unordered and of equal Cladistic analysis weight, MulTrees option in effect. Bootstrap values for For the analysis, three Protohermes species, P. infectus clades were calculated in 1000 replicates using a general (McLachlan, 1869), P. changninganus Yang and Yang, heuristic search, with branches with bootstrap values 1988, and P. niger Yang and Yang, 1988, were selected <50% collapsed. Bremer’s decay index was calculated as the outgroups. This is because P. infectus and using Autodecay version 4.0 (Eriksson 1998) and PAUP* P. changninganus are considered to have a basal status version 4.0b10. in Protohermes. Also, P. niger is considered to be the most primitive species of the P. costalis species-group. TAXONOMY Nine adult morphological characters were numeri- cally coded for all three species of the P. xanthodes Genus Protohermes van der Weele species-group and the three outgroups, which are listed Protohermes van der Weele (1907): 243. Type species below. All the characters presently selected were Hermes anticus Walker (1853): 205, original obtained from the male genitalia except the first two, designation. which were obtained from the head and wings. The data Allohermes Lestage (1927): 100. Type species Protoher- matrix is given in Table 1. The character state coding is: mes davidi van der Weele (1909): 254, original 0, plesiomorphic; 1–3, apomorphic. designation. (Synonymized by Glorioso 1981.) 1 Lateral ocelli: close to median ocellus, 0; widely apart from median ocellus, 1. General characters. Body medium to large sized 2 Forewing with costal areas: hyaline, 0; dark or (forewing length 28–65 mm). Body pale yellow to yel- marked by several dark stripes, 1. lowish brown, sometimes blackish brown. 3 Male ninth sternum: narrower than ninth tergum, 0; Diagnosis. Head robust, postocular spines present or wider than ninth tergum, 1. absent; vertex often with several dark marks. Posterior 4 Male ninth sternum with median portion: not pair of ocelli close to or widely apart from median inflated, 0; distinctly inflated, 1. ocellus, median ocellus usually flattened and transverse. 5 Male tenth tergite: simple digitiform, 0; long band- Antenna subserrate, approximately as long as the like, 1; subcylindrical, 2; claviform, 3. head plus prothorax. Clypeal margin entire. Labrum subtriangular. Table 1 Character matrix for three species of the Prothorax longer than wide, pronotum with various Protohermes xanthodes species-group and three outgroups patterns of dark marks. Wings slightly smoky brown to blackish brown, often with several yellowish or whitish Characters marks, showing distinct specific variations. Rs 8–11- Taxon 1 23456789 branched, last branch bifurcate or trifurcate; 6–14 P. infectus 0 10000000 crossveins between R1 and Rs; M1+2 4–9-branched, M3+4 P. changninganus 0 10010000 2–4-branched; 1A three-branched. P. niger 1 10021000 Male ninth tergum often shorter than wide, with ante- P. grandis 1 01131010 rior margin incised in an arched manner; ninth tergum P. immaculatus 1 01131010 broad, often with V-shaped, arched, or trapezoidal pos- P. xanthodes 1 00031101 terior incision; tenth tergite slender digitiform, short 0, Plesiomorphic state; 1–3, apomorphic state. claviform, short subcylindrical, flattened valvate, or

400 Entomological Science (2006) 9, 399–409 © 2006 The Entomological Society of Japan Systematics of the P. xanthodes species-group elongated band-like; ninth gonostylus unguiform; tenth Protohermes grandis (Thunberg) (Figs 1,4–10) sternite often arched, with pair of lateral lobes digiti- Hemerobius grandis Thunberg (1781): 28. form or tubercle. Female eighth sternum strongly sclerotized, often produced posteriorly; ninth gonocoxite Diagnosis. Head with three pairs of black marks later- membranous, ﬂattened; ninth gonostylus short and dig- ally on vertex; pronotum with two pairs of black vittae itiform, articulated with gonocoxite; tenth tergite often on each side; male tenth tergite short, claviform, with incised from side by cerci, forming one dorsal and one apex produced into two processes, and inner process ventral lobe. bearing one setal tuft. Male. Body length 24–43 mm; forewing length 33– 51 mm, hindwing length 29–46 mm. Protohermes xanthodes species-group Head yellowish brown, with two pairs of large sub- Diagnostic characters. Head and pronotum yellow to quadrate and one pair of small suboval black marks on yellowish brown, with several blackish marks. Ocelli lateral portion of vertex; postocular spine absent. Com- widely apart. Wing slightly smoky brown, usually with pound eyes brown, ocelli yellow with black inner margin. yellowish marks; costal areas without dark stripes. Male Median ocellus rather ﬂattened and transverse; posterior tenth tergite short, claviform, with one tuft on inner pair widely apart, distance between them about twice subdistal portion. Male tenth sternite wide, with elon- the width of median ocellus but shorter than that gate lateral lobes. Female genitalia simple, without sac- between antennal sockets. Antenna black, scape and like lobes on ninth abdominal segment. pedicel yellowish brown. Mouthparts yellowish brown; Remarks. This species-group appears to be closely maxillary and labial palpi with distal two segments infus- related to the P. costalis species-group by having similar cate; mandible with distal half blackish brown. widely separated ocelli and the male tenth tergite bear- Prothorax yellowish brown; pronotum with two pairs ing a tuft of setae, but it can be easily separated from of narrow black vittae near lateral margins, somewhat the latter by the claviform male tenth tergite and the connected at midlength. Meso- and metathorax yellow- simple female genitalia without lateral lobes. In the ish brown, dorsally with lateral portion brown. Tho- P. costalis species-group, the male tenth tergite is subcy- racic pilosity yellow, much longer on meso- and lindrical with a more or less incised tip, and the female metathorax. Legs yellowish brown with short, dense, genitalia bear a pair of sac-like lateral lobes on the ninth yellowish setae; apices of tibiae wide, grayish brown, abdominal segment. tarsi grayish brown, tarsal claws reddish brown. Forew- ings pale smoky brown, with pair of large and small yellowish marks at base, one large and three small yel- Key to species of the P. xanthodes group lowish marks at middle, and one round yellowish mark 1 Male ninth gonostylus thick (Fig. 19); male tenth ster- at apical 1/3. Hindwings pale smoky brown except basal nite moderately sclerotized, simple, with spinous tip half hyaline, with round yellowish mark at middle and (Fig. 21); female eighth sternum with posterior mar- at apical 1/3, respectively. Veins brown, with veins much gin feebly produced (Fig. 22).....P. xanthodes Navás paler in anal areas and yellowish marks. Rs ten- or 11- – Male ninth gonostylus slender (Figs 7,14); male branched, last branch bifurcate; eight or nine crossveins

tenth sternite feebly sclerotized, median portion between R1 and Rs; M1+2 ﬁve- or six-branched, M3+4 enlarged, with tip rounded (Figs 8,15); female eighth two-branched; 1A three-branched. sternum with posterior margin strongly produced Abdomen brown with venter and genitalia yellow. (Figs 9,16)...... 2 Ninth tergum (Fig. 5) transversely widened, with arched 2 Large-sized (forewing length 33–65 mm); wings with anterior and posterior margins. Ninth sternum (Fig. 6) several distinct yellowish marks; male tenth tergite much broader than ninth tergum; posterior margin with apex produced into two processes (Fig. 5); deeply incised, trapezoidal, forming pair of acutely pro- female eighth sternum with posterior margin medially duced processes; central portion apparently inﬂated. incised (Fig. 10) ...... P. grandis (Thunberg) Ninth gonostylus (Fig. 7) slender, unguiform, slightly – Small-sized (forewing length 26–39 mm); wings with curved dorsad. Tenth tergite (Figs 5,6) short, claviform, small indistinct yellowish marks; male tenth tergite with apex produced into two processes, its inner process with simple apex (Fig. 12); female eighth sternum bearing one setal tuft. Tenth sternite (Fig. 8) arched, with posterior margin truncate (Fig. 17) ...... medially elevated at anterior and posterior margins; lat- ...... P. immaculatus Kuwayama eral lobes elongated, digitiform, slightly incurved.

Figures 1–3 Adults of Protohermes spp. 1 Protohermes grandis (Thunberg) (male). 2 Protohermes immaculatus Kuwayama (male). 3 Protohermes xan- 3 thodes Navás (male from Korea). Scale lines: 5.0 mm.

Female. Body length 25–45 mm; forewing length 42– ventral view, with posterior margin slightly incised, 65 mm, hindwing length 38–53 mm. Color similar to forming pair of roundly produced processes; ninth male. gonocoxite broad, ventro-distal portion concave, with Female eighth sternum (Figs 9,10) strongly sclero- small digitiform processes at tip; tenth tergite short, tized, subtriangular in lateral view, subtrapezoidal in with posterior margin triangularly incised from side,

Figures 4–10 Protohermes grandis (Thunberg). 4 Male head and prothorax, dorsal view; 5 male genitalia, dorsal view; 6 male genitalia, ventral view; 7 male ninth gonostylus, caudal view; 8 male tenth sternite, ventral view; 9 female genitalia, lateral view; 10 female eighth sternum, ventral view. Scale lines: 1.0 mm. leaving one subtriangular dorsal lobe and one semicir- kawa, Chiba Pref., 29.v.1987, K. Fukuyama; 1 2 cular ventral lobe. (HC), Ino-kawa, Chiba Pref., 17, 29.viii.1989, K. Fuku- Specimens examined. Japan: 1 3 (HC), Akan- yama; 1 1 (HC), Yozawa-gawa, Itsukaichi, Tokyo, gawa, Akan, Hokkaido Prefecture (Pref.), 5.viii.1990, 13.vii.1990, S. Uchida; 1 (HC), Yozawa-gawa, Itsuka- R. Kuranishi; 1 2 (HC), Akabira, Hokkaido Pref., ichi, Tokyo, 2.vii.1990, S. Uchida; 1 (HC), Todori- 20.vii and 1.viii.1996, S. Hara; 1 (HC), Uryu, Hok- machi, Hachioji, Tokyo, 1.vii.2001, F. Hayashi; 1 kaido Pref., 1.viii.1996, S. Hara; 1 (HC), Syakotan, (HC), Setagaya-ku, Tokyo, 21.v.1951, N. Iwasaki; Hokkaido Pref. 1997, reared from larva, F. Hayashi; 1 13 6 (HC), Morito-gawa, Miura Peninsular, (HC), Oguni-gawa, Iwate Pref., 16.viii.1983, S. Uchida; Knagawa Pref., F. Hayashi; 3 3 (HC), Yataro- 4 2 (HC), Natori-gawa, Miyagi Pref., reared gawa, Susugaya, Kanagawa Pref., F. Hayashi; 1 from larvae, F. Hayashi; 1 (HC), Aizu-tajima, Fuku- (CAU), Matsuda-machi, Kanagawa Pref., 7.vi.1997, S. shima Pref., 22.vii.1986, G. Mori; 3 1 (HC), Hara; 1 (HC), Doshi-mura, Tsuru-gun, Yamanashi Irihirose-mura, Kitauonuma-gun, Niigata Pref., Pref., 31.vii.1993, K. Waki; 1 (HC), Uenohara, 27.vi.1998, M. Yamamoto; 4 (HC), Nasu-machi, Nirasaki, Yamanashi Pref., 28.vi.1994, K. Kariya; 1 Tochigi Pref., 28.vii.1996, K. Waki; 1 (HC), Ino- (HC), Hosaka-machi, Nirasaki, Yamanashi Pref.,

Figures 11–17 Protohermes immaculatus Kuwayama. 11 Male head and prothorax, dorsal view; 12 male genitalia, dorsal view; 13 male genitalia, ventral view; 14 male ninth gonostylus, caudal view; 15 male tenth sternite, ventral view; 16 female genitalia, lateral view; 17 female eighth sternum, ventral view. Scale lines: 1.0 mm.

7.vii.1995, K. Kariya; 1 (HC), Fujimi-kogen, Nagano gawa, Miyazaki Pref., reared from larvae, F. Hayashi; Pref., 14.vii.1988, K. Hatta; 1 (HC), Oppara, Kamo, 6 5 (HC), Sasu-gawa, Tsushima Island, Gifu Pref., 30.vii.1973, F. Kamiya; 1 (HC), Ohara, Nagasaki Pref., reared from larvae, F. Hayashi; 8 Ono-gun, Gifu Pref., 21.vi.1997, M. Yamamoto; 1 7 (HC), Shiratani-gawa, Yaku-shima Island, (HC), Miyawaki, Mikata-gun, Hyogo Pref., Kagoshima Pref., reared from larvae, F. Hayashi; 1 16.viii.1989, N. Yamakawa; 5 2 (1 in CAU, (SEMCAS), no detailed locality, 2.vi.1920. others in HC), Gunke-machi, Tottori Pref., 17.vii.1996, Distribution. Japan (Hokkaido, Okushiri Island, Hon- M. Yamamoto; 1 (HC), Nakamura-shi, Kochi Pref., shu, Sado Island, Oki-Togo Island, Shikoku, Kyushu, 1.vii.1995; 1 1 (HC), Ajimu-machi, Usa-gun, Oita Tsushima Island, Amakusa Islands, Yakushima Island, Pref., 9.viii.1984, T. Torii; 1 (HC), Jigoku, Aso-gun, Tanegashima Island; Hayashi 2005). Kumamoto Pref., 6.viii.1987, E. Matsui; 1 (HC), Remarks. This species is common in Japan and easily Mount Ichifusa, Kumamoto Pref., 31.vii.1989, E. Mat- distinguished by the large body size and the bright yel- sui; 1 (HC), Ue, Kuma-gun, Kumamoto Pref., lowish wing color pattern. It appears to be closely 23.vii.1987, E. Matsui; 4 5 (HC), Minami- related to P. immaculatus because the two species share

404 Entomological Science (2006) 9, 399–409 © 2006 The Entomological Society of Japan Systematics of the P. xanthodes species-group a slender male ninth gonostylus and a female eighth strongly curved dorsad. Tenth tergite (Figs 12,13) sim- sternum with the posterior margin strongly produced, ple, claviform, as long as ninth tergum, bearing inner but can be easily separated by the unique male tenth setal tuft on subdistal portion. Tenth sternite (Fig. 15) tergite with two processes at the tip. In P. immaculatus, arched, medially elevated at anterior and posterior mar- the male tenth tergite is simple without any additional gins, posterior margin with one small median incision; process (Kuwayama 1964). lateral lobes elongated, digitiform, directed dorsad and slightly incurved. Protohermes immaculatus Kuwayama Female. Body length 20–30 mm; forewing length 31– (Figs 2,11–17) 39 mm, hindwing length 28–35 mm. Color similar to male. Protohermes immaculatus Kuwayama (1964): 25. Female eighth sternum (Figs 16,17) strongly sclero- Diagnosis. Body small-sized; vertex with three pairs of tized, subtriangular in lateral view, subtrapezoidal in black marks; occiput with pair of black marks; mandi- ventral view, with truncate posterior margin; ninth ble mostly blackish brown; wings with small indistinct gonocoxite broad, ventro-distal portion concave, with yellowish marks; male tenth tergite simple, claviform, small digitiform processes at tip; tenth tergite short, with one inner setal tuft on subdistal portion. with posterior margin roundly incised from side, leaving Male. Body length 19–26 mm; forewing length 26– a thick subtriangular dorsal lobe and a semicircular 33 mm, hindwing length 24–29 mm. ventral lobe. Head yellow, vertex with three pairs of black marks Specimens examined. Japan: 2 (HC), Sumiyo, on lateral portion, marks of anterior pair large, sub- Amami-oshima Island, central Ryukyus, 22.vi.1987, K. quadrate, those of postero-outer pair subtriangular, and Uesugi; 5 6 (1 in CAU, others in HC), those of postero-inner pair smallest and suboval; pos- Kawauchi-gawa, Amami-oshima Island, reared from tocular spine present, feebly developed. Occiput with larvae, F. Hayashi; 4 7 (1 in CAU, others in pair of black marks on each side. Compound eyes black- HC), Asato-gawa, Amami-oshima Island, reared from ish brown, ocelli yellow with black inner margin. larvae, F. Hayashi; 2 1 (HC), Kametoku-gawa, Median ocellus rather ﬂattened and transverse; poste- Tokunoshima Island, central Ryukyus, reared from lar- rior pair widely apart, distance between them as long as vae, F. Hayashi; 1 1 (HC), Akirigami-gawa, Tokun- that between antennal sockets but less than twice the oshima Island, reared from larvae, F. Hayashi; 1 (HC), width of median ocellus. Antenna black, with scape and Mikyo, Tokunoshima Island, 8.vii.1989, H. Karube; pedicel yellow. Mouthparts yellow; galea, maxillary, and 4 1 (HC), Shirase-gawa, Kume-jima Island, cen- labial palpi much darker; mandible mostly blackish tral Ryukyus, reared from larvae, F. Hayashi. brown, with proximal inner portion much paler. Distribution. Japan (Amami-oshima Island, Tokun- Prothorax yellow; pronotum with pair of narrow oshima Island, Kume-jima Island). black vittae near each lateral margin. Meso- and met- Remarks. This species is the smallest known Protoher- athorax yellow, dorsally with lateral portion much mes species. As noted in the remarks for P. grandis darker. Thoracic pilosity yellow, much longer on meso- given above, P. immaculatus appears to be related to and metathorax. Legs yellow with short, dense, yellow- P. grandis, but can be easily distinguished because the ish setae; tarsi mostly grayish yellow, tarsal claws reddish wings have small and indistinct yellowish marks. brown. Wings pale smoky brown without any marks, forewing with small indistinct yellowish marks around Protohermes xanthodes Navás (Figs 3,18–23) stem of Cu1. Veins pale brown, with veins pale yellow Protohermes xanthodes Navás (1913): 427. in anal areas. Rs 9 to 11-branched, last branch bifurcate; Protohermes rubidus Stitz (1914): 201. syn. nov.

5–8 crossveins between R1 and Rs; M1+2 ﬁve- or six- Protohermes martynovae Vshivkova (1995): 24. syn. branched, M3+4 two-branched; 1A three-branched. nov. Abdomen blackish brown. Ninth tergum (Fig. 12) transversely widened, with arched anterior margin and Diagnosis. Body medium-sized; vertex with three pairs shallowly incised posterior margin. Ninth sternum of black marks; wings with yellowish marks; male tenth (Fig. 13) much broader than ninth tergum; posterior tergite simple, claviform, with inner median margin margin deeply incised, trapezoidal, forming pair of incised, bearing one inner setal tuft on subdistal portion; acutely produced processes; central portion apparently male ninth gonostylus strongly developed with one inﬂated. Ninth gonostylus (Fig. 14) slender, unguiform, apical claw.

Figures 18–23 Protohermes xanthodes Navás. 18 Male head and prothorax, dorsal view; 19 male genitalia, dorsal view; 20 male ninth sternum, ventral view; 21 male tenth sternite, ventral view; 22 female genitalia, lateral view; 23 female eighth sternum, ventral view. Scale lines: 1.0 mm.

Male. Body length 23–35 mm; forewing length 31– round, and yellowish mark at apical 1/3. Hindwings 42 mm, hindwing length 28–37 mm. pale smoky brown except basal half hyaline, with a Head yellow to yellowish brown, vertex with three round yellowish mark at the middle and apical 1/3, pairs of black marks on lateral portion, marks of ante- respectively. Veins pale brown, but pale yellow in yel- rior pair large and subquadrate, those of the postero- lowish marks and basal half of hindwing. Rs ten- outer pair subtriangular, and those of the postero-inner branched, last branch bifurcate or trifurcate; 11 cross- pair smaller and suboval; postocular spine present, fee- veins between R1 and Rs; M1+2 five- or six-branched, bly developed. Compound eyes blackish brown, ocelli M3+4 four-branched; 1A three-branched. yellow with black inner margin. Median ocellus flat- Abdomen yellow to brown. Ninth tergum (Fig. 19) tened and transverse; posterior pair widely apart, dis- nearly rectangular, with trapezoidal anterior margin and tance between them nearly as long as that between shallowly incised posterior margin. Ninth sternum antennal sockets and twice the width of median ocellus. (Fig. 20) much narrower than ninth tergum; posterior Antenna black, with scape and pedicel yellow. Mouth- margin deeply incised, trapezoidal, forming a pair of parts yellow; mandible with apical half blackish brown. acutely produced processes; central portion feebly Thorax yellow to yellowish brown; pronotum with a inflated. Ninth gonostylus (Fig. 19) thick, directed pair of black vittae near each margin; meso- and met- inward, with one slender distal claw. Tenth tergite anotum pale brown laterally. Thoracic pilosity yellow, (Fig. 19) simple claviform, slightly shorter than ninth much longer on meso- and metathorax. Legs yellow tergum, with inner median margin incised, bearing one with short, dense, yellowish setae; tibiae and tarsi pale inner setal tuft on subdistal portion. Tenth sternite brown, tarsal claws reddish brown. Forewings pale (Fig. 21) simple, arched; lateral lobes moderately sclero- smoky brown, with a large yellowish mark at base, three tized, digitiform, directed dorsally and slightly incurved, to four small yellowish marks at middle, and a small, with spinous apex.

Female. Body length 37–52 mm; forewing length 45– Yang; 1 1 (IZCAS), Jiangxi, Yiyang, 12/13.v.1975, 51 mm, hindwing length 40–47 mm. Color similar to Y. W. Zhang; 1 (IZCAS), Jiangxi, De-an, Pengshan- male. linchang, 13.v.1980; 7 5 (CAU), Zhejiang, Female eighth sternum (Figs 22,23) strongly sclero- Lishui, v.17/vi.25. 1982; 1 (CAU), Fujian, v.1990; 1 tized, subtrapezoidal in lateral view, suboval in ven- 4 (CAU), Guangdong, Wengyuan, v.1974, R. Z. Wu; tral view, with truncate posterior margin; ninth 1 (CAU), Guangxi, Guilin, 150 m, 8.vi.1984; 5 gonocoxite subtriangular, with small digitiform pro- 9 (CAU), Guangxi, Lingchuan, Lingtian, 275 m, 4/ cesses at tip; tenth tergite short, with posterior mar- 7.vi.1984, F. S. Li and D. Yang; 3 (CAU), Guangxi, gin roundly incised from side, leaving one thick Guilin, Yanshan, 7/8.v.1964; 2 (IZCAS), Guangxi, subtriangular dorsal lobe and one semicircular ven- Guilin, Yanshan, 13/16.v.1963, C. G. Wang; 1 tral lobe. (IZCAS), Guangxi, Jinxiu, Luoxiang, 200 m, 15.v.1999, Specimens examined. China: 1 (IZCAS), Liaoning, X. Z. Zhang; 1 (IZCAS), Guangxi, Xing-an, Maoers- Caohekou, 18.vii.1958; 1 (CAU), Shandong, Taian, han, 400 m, 12.vii.1985; 1 (CAU), Guizhou, Xishan, 20.vi.1962; 1 (CAU), Beijing, Pinggu, 1976; 3.vii.1964; 3 (CAU), Chongqing, Beibei, x.1955; 1 (CAU), Beijing, Campus of Beijing Agricultural Uni- 1 1 (CAU), Sichuan, Ya-an, 1973; 1 (CAU), versity, 1976; 1 (CAU), Beijing, Miyun, 9.vi.1960; 1 Sichuan, Ya-an, viii.1984; 1 (IZCAS), Sichuan, Peng- (CAU), Beijing, Miaofengshan, 24.vi.1955, L. S. Zhao; shui, 750 m, 9.vii.1989, L. L. Yang; 1 (IZCAS), 1 (CAU), Beijing, Xiangshan, 20.vi.1980, C. H. Yang; Sichuan, Pengshui, 750 m, 13.vii.1989, X. C. Zhang. 2 (CAU), Beijing, Dingjiatan, 28/30.vii.1960, Korea: 2 (1 in CAU, 1 in HC), Yongsuchon River, F. S. Li; 6 (IZCAS), Beijing, Sanpu, 25.vii.1967, Yongsuri, Changaum, Kijiang-gun, reared from larvae, J. M. Zhao; 1 (IZCAS), Beijing, Badaling, 700 m, F. Hayashi. 25.vi.1964, C. B. Liao; 1 (IZCAS), Beijing, Chang- Distribution. China (Anhui, Beijing, Gansu, Guang- ping, 9.vii.1959; 3 (IZCAS), Beijing, Sanpu, 22/ dong, Guangxi, Guizhou, Hebei, Henan, Hubei, Hunan, 24.vii.1964, C. B. Liao; 1 (IZCAS), Beijing, Sanpu, Jiangxi, Liaoning, Shandong, Shanxi, Shaanxi, Sichuan, 9.vii.1964, Q. Zhou; 1 (IZCAS), Beijing, Sanpu, Yunnan, Zhejiang); Korea (Kijiang-gun); Russia (south 5.viii.1964, Q. Zhou; 1 (CAU), Beijing, Fangshan, Primorskij). Hebeigongshe, 6.vi.1960, C. K. Yang; 1 (CAU), Remarks. This species was described by Navás (1913) Beijing, Miaofengshan, 10.viii.1964, X. M. Cai; 1 1 based on one female specimen from Yunnan (Dali), (CAU), Shanxi, Daning, Xifeng, 1980; 5 (CAU), with only one figure of the pronotum in his original Shanxi, Taigu, 26/30.vii.1964, C. H. Yang; 1 (CAU), description, which has been difficult to use in identifi- Gansu, Wenxian, Gaoloushan 1700 m, 7.viii.1960, C. cation. However, this species’ identity was clarified K. Yang; 1 (CAU), Gansu, Dianbu, Lazikou, 1700 m, after the redescriptions by Tjeder (1937, 1954). Cur- 12.viii.1980, C. K. Yang; 1 3 (IZCAS), Gansu, rently, P. xanthodes is the most common of the Proto- Chengxian, Feilongxia, 1020 m, 4.vii.1999, T. L. He; hermes xanthodes species-group species in China, 1 (CAU), Shannxi, Ankang, 10.vii.1981; 1 (CAU), with a wide distribution. Protohermes xanthodes is Shannxi, Yan-an, viii.1981; 1 (CAU), Shannxi, Han- herein recorded from Korea for the first time, with zhong, v.1981; 1 (CAU), Shannxi, Y. Zhou; 1 specimens that appear paler and smaller than those (IZCAS), Shannxi, Foping, 900 m, 27.vi.1999, J. Yao; from China. 1 7 (IZCAS), Shannxi, Foping, 890 m, Protohermes rubidus was established by Stitz in 1914 26.vi.1999, T. L. He; 2 (IZCAS), Shannxi, Foping, based on specimens from China (Qingdao, Shandong). 900 m, 27.vi.1999, C. D. Zhu; 1 3 (CAU), Henan, Banks (1940) mentioned that this species might be the Xishan, Huangshian, 1000 m, 16.vii.1998, X. C. Shen same as P. xanthodes. In the present paper, in accor- and Y. D. Ren; 2 (CAU), Henan, Huanglianshu, dance with the original illustrations of the male genitalia 1600 m, 6.vi.2000, X. C. Shen and Y. D. Ren; 9 of P. rubidus, which are the same as those of 16 (CAU), Henan, Shangcheng, Huangbaishan, P. xanthodes, we have placed P. rubidus as a junior syn- 14.vii.1999, X. C. Shen and Y. D. Ren; 1 (IZCAS), onym of P. xanthodes. Anhui, Shitai, 1.vi.1975; 1 (CAU), Hubei, Wuxia, According to the original illustrations of the male Guandukou, 8.vi.1951, B. L. Lu; 1 (CAU), Hubei, genitalia, P. martynovae Vshivkova, 1995, seems to be Shennongjia, Songbai, 22.vi.1977, S. L. Liu; 2 2 within the range of variation of P. xanthodes. For this (IZCAS), Hunan, Dong-an, 3/10.v.1954; 1 (CAU), reason, this species is considered to be one of the north- Hunan; 1 (CAU), Jiangxi, Shangrao, 30.iv.1973, C. K. eastern populations of P. xanthodes.

Figure 24 The most parsimonious tree for all species of the Protohermes xanthodes species-group (L = 11, CI = 1.00, RI = 1.00). Numbers below squares correspond to characters as described in the text: numbers above squares correspond to states. Bremer’s decay index and bootstrap values >50% (1000 replicates) are indicated in the circles at the nodes (Bremer’s decay index on the left side of the diagonal line, bootstrap values on the right side of the diagonal line).

Figure 25 Distribution map of the Protohermes xanthodes species-group. () and ( ), P. xanthodes; ( ), P. grandis; circled area, P. immaculatus.

RESULTS AND DISCUSSION acter 6: 1). Within the P. xanthodes group, the sister relationship between P. grandis and P. immaculatus is A heuristic search resulted in only one most parsimoni- supported by a broader male ninth sternum with ous tree (length (L) = 11, consistency index (CI) = 1.00, strongly inﬂated median portion (characters 3: 1, 4: 1) retention index (RI) = 1.00). The most parsimonious and a highly modiﬁed male tenth sternite with the tree is shown in Figure 24; Bremer’s decay index and median portion enlarged (character 8: 1). The thick bootstrapping values over 50% are also represented. male ninth gonostylus (character 7: 1) and the moder- The P. xanthodes species-group formed a monophyl- ately sclerotized lateral lobes of the male tenth sternite etic group supported by immaculate costal areas of (character 9: 1) represent the autapomorphic characters forewings (character 2: 0) and a claviform male tenth of P. xanthodes. tergite (character 5: 3). Its sister relationship with the Considering the distributions of the P. xanthodes spe- P. costalis species-group is supported by three synapo- cies-group (Fig. 25), P. xanthodes is widely distributed morphic characters: the widely separated ocelli (charac- in China, extending northward to the Russian Far East ter 1:1), a subcylindrical male tenth tergite (character 5: and Korea. Protohermes grandis is restricted to Palae- 2), and a male tenth tergite bearing one setal tuft (char- arctic Japan, and P. immaculatus is restricted to only

some islands of the central Ryukyus, which belong to Kimmins DE (1948) Notes on the genus Protohermes Weele southern Japan. Both these species are recognized to be (Megaloptera) with description of two new species. distributed along the West Pacific Island Arc, which Annals and Magazine of Natural History 1, 765–781. might have been separated from the Asian mainland Kuwayama S (1964) On the Neuroptera from Amami-oshima during the transgression of the Pacific Ocean after the and Yakushima. Mushi 38, 25–31. Lestage JA (1927) La fauna entomologique Indochinoise, 2: mid-Pleistocene (Zhang 1999). Therefore, speciation of les Megalopteres. Bulletin et Annales de la Société Royale P. grandis and P. immaculatus might take place due to d’Entomologie de Belgique 67, 71–90, 93–119. the vicariance between the Asian mainland and the West Liu XY, Yang D (2005) Revision of the Protohermes chang- Pacific Island Arc. ningensis species-group from China (Megaloptera: Cory- dalidae: Corydalinae). Aquatic Insects 27, 167–178. ACKNOWLEDGMENTS Liu XY, Yang D (2006a) Revision of the Protohermes species from Tibet, China (Megaloptera: Corydalidae). Zootaxa We are grateful to Dr Xianwei Liu (Shanghai) and Dr 1199, 49–60. Weinian Zhang (Shanghai) for the loan of several spec- Liu XY, Yang D (2006b) Systematics of the Protohermes imens deposited in the Entomological Museum of davidi species-group (Megaloptera: Corydalidae), with Shanghai, Chinese Academy of Science, and to several notes on its phylogeny and biogeography. Invertebrate Systematics 20, 477–488. Japanese colleagues for providing useful materials. This Penny, DE (1993) The phylogenetic position of Chloroniella research was supported by the National Natural Science peringueyi (Megaloptera: Corydalidae) and its zoogeo- Foundation of China (30370174, 30225009). graphic significance. Entomological News 104, 17–30. Navás L (1913) Neuroptera Asiatica. II series. Revue Russe REFERENCES D’entomologie 13, 424–430. Swofford DL (2002) PAUP*: Phylogenetic Analysis Using Par- Banks N (1940) Report on certain groups of Neuropteroid simony (*and Other Methods), Version 4.0b10. Sinauer insects from Szechwan, China. Proceedings of the United Associates, Sunderland, MA, USA. States National Museum 88, 173–220. Tjeder B (1937) Schwedisch-Chinesische wissenschaftliche Contreras-Ramos A (1998) Systematics of the Dobsonfly Expedition nach den nordwestlichen Provinzen Chinas. Genus Corydalus Latreille (Megaloptera: Corydalidae). 62. Neuroptera. Arkiv for Zoologi 29A, 1–36. Thomas Say Monographs, Entomological Society of Tjeder B (1954) Genital structures and terminology in the America, Maryland, USA. order Neuroptera. Entomologische Meddelelser 27, 23– Eriksson T (1998) Autodecay, Version 4.0. (Program distrib- 40. uted by the author.) Bergius Foundation, Royal Swedish Vshivkova TS (1995) Megaloptera. In: Kozlov MA, Makarch- Academy of Sciences, Stockholm, Sweden. enko EA (eds) Keys to the Insects of the Russian Far East, Glorioso MJ (1981) Systematics of the dobsonfly subfamily pp. 9–34. Nauka, St. Petersburg, Russia. (In Russian.) Corydalinae (Megaloptera: Corydalidae). Systematic Ento- van der Weele HW (1907) Notizen uber Sialiden und Beschrei- mology 6, 253–290. bung einiger neuen Arten. Notes from the Leyden Hayashi F (1989a) Protohermes immaculatus Kuwayama Museum 28, 227–264. (Megaloptera, Corydalidae). Nature and Insect 24, 19– van der Weele HW (1910) Megaloptera: monographic revi- 21. (In Japanese.) sion. Collections Zoologiques Du Baron Edm. De Selys Hayashi F (1989b) Discovery of the adult dobsonfly Protoher- Longchamps Fasc 5, 1–93. mes sp. (Tanida, 1974) on Ishigaki and Iriomote Islands. Yang D (1985) Contribution to the taxonomy of Corydalinae The Insectarium 26, 354–356. (In Japanese.) from China (Insecta: Megaloptera). MS thesis, Beijing Hayashi F (1990) Notes on Protohermes grandis. Iden 44, 72– Agricultural University, Beijing, China. (In Chinese.) 77. (In Japanese.) Yang CK, Yang D (1988) New species of Corydalinae from Hayashi F (2005) Megaloptera. In: Kawai T, Tanida K (eds) Yunnan (Megaloptera: Corydalidae). Zoological Research Aquatic Insects of Japan: Manual with Keys and Illustra- 9, 45–60. tions, pp. 379–386. Tokai University Press, Kanagawa, Zhang YZ (1999) Zoogeography of China. Science Press, Japan. (In Japanese.) Beijing, China. (In Chinese.)