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. © 2006 The Entomological Society of Japan X. Y. Liu et al. 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 scle- rotized, 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