Zootaxa, Systematics of the Protohermes Costalis Species-Group

Systematics of the Protohermes costalis species-group (Megaloptera: Corydalidae)

XINGYUE LIU1, FUMIO HAYASHI2 & DING YANG1,* 1Department of Entomology, China Agricultural University, Beijing 100094, China. E-mail: [email protected] or [email protected] 2Department of Biology, Tokyo Metropolitan University, Minamiosawa 1–1, Hachioji, Tokyo 192–0397, Japan. *Corresponding author

Table of contents

Abstract ...... 2 Introduction ...... 2 Material and methods...... 2 Results ...... 7 Discussion ...... 7 Systematics...... 11 Genus Protohermes Weele 1907 ...... 11 The Protohermes costalis species-group ...... 11 Key to species of the P. c o st ali s species-group 12 Protohermes acutatus sp. nov...... 13 Protohermes arunachalensis Ghosh ...... 16 Protohermes basiflavus Yang ...... 18 Protohermes basimaculatus sp. nov...... 19 Protohermes costalis (Walker) ...... 21 Protohermes disjunctus sp. nov...... 23 Protohermes fujianensis Yang & Yang ...... 25 Protohermes gutianensis Yang & Yang ...... 26 Protohermes hunanensis Yang & Yang ...... 28 Protohermes lii sp. nov...... 30 Protohermes niger Yang & Yang ...... 31 Protohermes orientalis sp. nov...... 33 Protohermes similis Yang & Yang ...... 35 Protohermes sinensis Yang & Yang ...... 36 Protohermes stigmosus sp. nov...... 38 Protohermes triangulatus sp. nov...... 39 Protohermes yangi sp. nov...... 41 Protohermes yunnanensis Yang & Yang ...... 43 Acknowledgements ...... 44 References ...... 45

Accepted by R.E. DeWalt: 9 Feb. 2007; published: 2 Apr. 2007 1 Abstract

The dobsonfly genus Protohermes is the largest genus in Megaloptera, with the richest species diversity and the widest geographic distributions. In the present study, the Protohermes costalis species-group is recognised, including 18 species. Eight species are described herein as new: Protohermes acutatus, Protohermes basimaculatus, Protohermes disjunctus, Protohermes lii, Protohermes orientalis, Protohermes stigmosus, Protohermes triangulatus and Protohermes yangi spp. nov. The interspecific phylogeny within the P. costalis species-group is reconstructed by a cladistic analysis. Discussions on the phylogeny and the biogeography of the P. costalis species-group are also made.

Key words: Corydalidae, Protohermes, new species, phylogeny, biogeography, Asia

Introduction

The megalopeteran genus Protohermes Weele is the largest genus of Corydalidae, with nearly 50 described species, which are distributed from far eastern to southeastern Asia. The adult is medium to large sized, usually with yellowish body coloration and several yellowish or whitish markings on wings. The male genitalia show various modifications and provide useful evidence for specific identification. The male tenth tergum varies considerably from elongated band-like structures to short subcylindrical processes. The male ninth sternum is modified to form a subgenital plate, with a posterior incision varying from deep to shallow and from V-shaped to trapezoidal. The female genitalia sometimes show a few interspecific differences in morphology. As with many corydalid species, Protohermes larvae are predaceous, living in clean streams of mountainous and wooded areas. The adults emerge from late spring to early autumn, are usually found near stream banks, and are readily collected at lights. The detailed natural history of Protohermes is well known only in Protohermes costalis (Walker), Protohermes immaculatus Kuwayama, and Protohermes grandis (Thunberg), from the studies by Hayashi (1989a, 1989c, 1990). The genus Protohermes was established by Weele (1907) and revised by many authors (Weele, 1910; Lestage, 1927; Kimmins, 1948; Glorioso, 1981; Yang, 1985). The genus is monophyletic, with Neurhermes as its sister. Together they are a sister lineage to the remaining genera within Corydalinae under Penny’s assumption (1993) or sister lineage to the rest of Corydalinae genera, excepting Chloroniella, with the most primitive phylogenetic status under Contreras-Ramos’ (1998) assumption. However, the interspecific phylogeny of the Protohermes species remains poorly studied so far. Recently, Liu & Yang (2005) have split the genus into several species groups. Protohermes costalis (Walker) is a common species in Oriental China. It is characterized by widely separated ocelli and a subcylindrical male tenth tergum bearing a ventral tuft. During this study, more species were found to be closely related to P. c o st al is . Thus, in the present paper, the Protohermes costalis species-group is recognized. All 18 group members, including eight species new to science, are keyed, described, and illus- trated. A preliminary phylogenetic analysis of the P. c o st a li s species group is undertaken based on morphological data. Geographic distributions are updated, and phylogenetic relationships and biogeography for the species group are also discussed.

Material and methods

Taxa studied Specimens were obtained from the following collections: the Entomological Museum of China Agricul- tural University (CAU), Beijing; Hebei University (HBU), Baoding; Institute of Zoology, Chinese Academy of Sciences (IZCAS), Beijing; Kunming Institute of Zoology, Chinese Academy of Sciences (KIZCAS), Kun-

2 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. ming; the Shanghai Entomological Museum, Chinese Academy of Sciences (SEMCAS), Shanghai; Shanghai Normal University (SNU), Shanghai; and Fumio Hayashi’s personal collection (HC), Tokyo, Japan. Genitalic preparations were made by macerating the apex of the abdomen in cold 10% KOH for 8–10 hours. The abdomen was transferred to glycerin for further examination and moved to a microvial with fresh glycerin, pinned below the specimen. Habitus photos of adults were obtained using a Nikon Coolpix 4500 digital camera. Drawings were made under the light microscope. Terminology generally follows that of Con- treras-Ramos (2004).

Cladistic analysis The phylogenetic analysis includes 35 mophological characters (Table 1) obtained from all 18 species of the P. c o st al is group. The cladogram was rooted using the following two Protohermes species, Protohermes infectus (McLachlan) and Protohermes xanthodes Navás. Although the phylogenetic relationships among all the Protohermes species have not been worked out, Protohermes infectus is considered herein to be a species possessing relatively more plesiomorphic characters than the other Protohermes species, such as the closely gathered ocelli, the male ninth gonostylus not eveloped by the ninth tergum, and the simple digitiformed male tenth tergum. Protohermes xanthodes seems to have the closest affinity with the P. c o st al is group in having similar widely separated ocelli and tuft on the male tenth tergum (Liu, Yang & Hayashi, 2006). Cladistic analysis was performed using PAUP* version 4.0b10 (Swofford, 2002) by using a heuristic parsimony analysis, with 100 random stepwise additions of taxa (TBR branch swapping) under ACCTRAN opti- mization, characters unordered and of equal weight, MulTrees option in effect. A branch and bound searching was also used. Bremer’s decay index was calculated with Autodecay version 4.0 (Eriksson, 1998) and PAUP* version 4.0b10. Bootstrap values for clades were calculated in 1000 replicates using a general heuristic search, the branches with bootstrap values <50% collapsed. Unambiguous characters were mapped using MacClade version 4.0 (Maddison & Maddison, 2000). Characters and character states employed in this analysis are listed below. Character states were scored 0–2 or ? (0 = plesiomorphic state, 1–2 = apomorphic state, ? = state unknown). 1. Head with post-ocular spine: (0) Present; (1) Absent. The post-ocular spine is present in many genera of Corydalinae. Most Protohermes species have a pair of post-ocular spines, though they are much weaker than that in Acanthacorydalis, Neoneuromus, etc. In most species of the P. c os ta l is group the post-ocular spine is absent except in P. basimaculatus sp. nov., P. niger Yang & Yang, and P. stigmosus sp. nov. with obtuse post-ocular spines (Figs. 45, 95, 123). 2. Posterior ocelli: (0) Close to each other; (1) Widely separated from each other. In P. xanthodes and all species of the P. c os ta l is group, the ocelli are distinctly widely separated, mostly with the distance between the posterior ocelli as long as that between the antennal bases. 3. Markings on vertex: (0) Present; (1) Absent. In P. basiflavus Yang, P. costalis, P. disjunctus sp. nov., P. lii sp. nov., P. stigmosus sp. nov., and P. triangulatus sp. nov., the head is pale yellow to yellowish brown without any markings on vertex (Figs. 38, 53, 60, 90, 123, 142). 4. Pronotum: (0) With two markings or more on each side; (1) With one vitta on each side; (2) Entirely black. In most P. costalis group members, there are at least two markings on each side, while there is only one vitta in P. fujianensis Yang & Yang, P. gutianensis Yang & Yang, P. hunanensis Yang & Yang, P. sinensis Yang & Yang, and P. y a ng i sp. nov., and the pronotum is entirely black in P. ni g er. 5. Forewing with round markings on apical 1/3: (0) With two radial veins at most through them; (1) With three or four radial veins through them. The round marking on forewing is an apomorphic character in many Protohermes and all Neurhermes species. The normal form of the apical markings is round with only two radial veins through them. However, in P. fujianensis, P. hunanensis, and P. yangi, the round markings are enlarged, with three or four radial veins through them (Figs. 12, 14, 22).

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 3 0 0 1 0 1 0 1 0 0 0 0 ? ? 0 0 0 ? 0 0 1 0 0 1 1 1 0 1 1 1 1 1 ? ? 1 1 1 ? 1 1 1 0 0 1 0 1 0 1 1 0 0 0 ? ? 1 1 1 ? 0 0 1 0 0 1 1 1 1 1 1 1 1 1 ? ? 1 1 1 ? 1 1 1 0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1 3 012345 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 0 1 0 1 1 0 0 0 1 0 0 0 0 2 0 0 1 0 0 1 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 0 1 1 1 0 0 1 1 1 0 1 1 1 1 1 1 0 1 1 1 1 1 1 1 . 0 0 1 1 1 0 1 1 1 1 2 1 0 1 1 0 0 1 2 1 0 0 1 1 1 0 1 1 1 1 1 1 0 1 1 0 0 1 1 1 2 0123456789 0 0 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 0 1 0 1 0 0 1 0 0 1 0 0 0 0 1 0 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 0 1 1 0 1 0 1 0 0 0 0 0 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 group used in the cladistic analysis used in the cladistic group 0 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 1 1 1 1 1 1 1 0 1 0 0 1 1 1 0 0 1 0 1 0 1 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0123456789 0 1 1 1 1 0 0 1 1 1 1 1 0 1 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 0 1 1 0 Protohermes costalis Protohermes 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 0 1 1 1 1 1 1 0 1 1 1 0 0 1 1 1 0 1 1 1 2 1 1 1 1 1 1 0 1 1 0 1 1 1 1 0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 1 1 1 0 2 0 0 1 0 0 1 0 0 0 0 0 1 0 1 1 0 0 0 1 0 0 0 0 1 0 0 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 1 1 1 0 1 1 1 1 1 1 0 1 1 1 0 1 1 1 Character state distribution for the for the distribution state Character TABLE 1. TABLE Characters Taxa 1 infectus P. 2 3 xanthodes P. 4 acutatus P. 5 arunachalensis P. 6 basiflavus 7 P. 8 basimaculatus P. 9 costalis P. disjunctus P. fujianensis P. gutianensis P. hunanensis P. lii P. niger P. orientalis P. similis P. sinensis P. stigmosus P. triangulatus P. yangi P. yunnanensis P.

4 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. 6. Forewing with stripes on costal areas: (0) Absent; (1) Present in most cellules; (2) Present in only basal 1–3 cellules. The immaculate forewing costal areas is apomorphic, while there are numerous dark stripes in the forewing costal cellules in many Protohermes species. However, in some Protohermes species, the dark stripes are present in only basal 1–3 costal cellules, such as P. basimaculatus.

7. Branches of M3+4: (0) Two branches; (1) More than three branches. Two branched M3+4 is a plesiomorphic character in most Protohermes species, while it is three or four branched in many species of the P. costalis group, which is considered to be apomorphic. 8. Male ninth sternum, posterior margin: (0) Incised;(1) Truncate. Penny (1993) notes that the posteriorly incised male ninth sternum is an apomorphic character in Protohermes, but in P. disjunctus and P. yunnanensis Yang & Yang, it is almost truncate (Figs. 62, 144). 9. Male ninth sternum, posterior incision: (0) Trapezoidal; (1) V-shaped. In the outgroups and most species of the P. c os ta l is group, the posterior incision of the male ninth sternum is trapezoidal except for P. basimaculatus, P. si ne n si s, P. yangi, and P. yunnanensis, which have a V-shaped incision (Figs. 46, 117, 130, 137). In P. gutianensis, the posterior incision is suboblong (Fig. 77) and considered to be the same state as the trapezoidal posterior incision in the present analysis. 10. Male ninth sternum, lateral margin: (0) Straight; (1) Curved. In P. i n fe ct u s, P. basimaculatus, P. c o st a - lis, P. niger, P. s in e ns is , P. s ti g m o su s, P. triangulatus, and P. yangi, the lateral margin of the male ninth sternum is straight (Figs. 46, 55, 97, 117, 125, 130, 137), while in P. xanthodes and the other species of the P. costalis group, the lateral margin of the male ninth sternum is more or less curved inward. 11. Male ninth sternum: (0) Narrower than ninth tergum; (1) Wider than ninth tergum. In P. acutatus sp. nov., P. basiflavus, P. c os ta l is , and P. disjunctus, the male ninth sternum is much wider than the ninth tergum (Figs. 26, 40, 55, 62), while the ninth sternum is much narrower than the male tergum in the outgroups and the other species of the P. costalis group. 12. Male ninth sternum, posterior processes: (0) Widely separated; (1) Close to each other. The posterior process of the male ninth sternum is formed by the posterior incision of the sternum. In P. infectus, P. basiflavus, P. basimaculatus, P. orientalis sp. nov., P. sinensis, and P. st ig m o s us , the posterior processes are widely separated (Figs. 40, 46, 103, 117, 125), while in P. xanthodes and the other species of the P. c os ta l is group, the processes are close to each other. 13. Male ninth sternum, length from base to tip of posterior process: (0) As long as ninth tergum; (1) As long as ninth plus tenth tergum. In P. acutatus and P. c os ta l is , the male ninth sternum is long, with its length from the base to the tip of the posterior process as long as the ninth plus tenth tergum. 14. Male ninth gonostylus: (0) Visible in dorsal view; (1) Invisible in dorsal view. In outgroups, the male ninth gonostylus is well developed and visible in dorsal view. In the P. costalis group, the male ninth gonostylus is enveloped by the ninth tergum and invisible in dorsal view. 15. Male ninth gonostylus: (0) Slightly curved; (1) Strongly curved. In P. basimaculatus, P. disjunctus, P. fujianensis, P. hunanensis, P. n i ge r, and P. yangi, the male ninth gonostylus is strongly curved inwards and dorsad (Figs. 49, 63, 70, 85, 98, 138), while the male ninth gonostylus is slightly curved in the outgroups and in the other species of the P. costalis group. 16. Male tenth tergum: (0) Not cylindrical; (1) Subcylindrical. The short subcylindrical male tenth tergum is synapomorphic to the P. costalis group. 17. Male tenth tergum, ventral process: (0) Feebly developed; (1) Well developed. The ventral process of the tenth tergum is an additional process located at the inner margin or ventral surface of the tenth tergum in some Protohermes species that often bears a tuft on the male tenth tergum. In P. acutatus, P. basiflavus, P. co s - talis, P. gutianensis, P. n i ge r, P. triangulatus, and P. yunnanensis, the ventral process is well developed, while in the other species of the P. costalis group, the process is feebly developed or even absent. 18. Male tenth tergum with tuft: (0) Absent; (1) Present. In P. xanthodes and all the species of the P. c o st a - lis group, there is one small tuft on the male tenth tergum (Fig. 26).

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 5 19. Male tenth tergum, position of tuft: (0) At inner margin of tenth tergum; (1) At ventral surface of tenth tergum. In P. xanthodes, the tuft is located at the inner margin of the male tenth tergum, whereas in all the species of the P. costalis group, the tuft is located at the ventral surface of the male tenth tergum. 20. Male tenth tergum: (0) Not incised at tip; (1) Incised at tip. In the outgroups and P. niger, the male tenth tergum is not incised at tip, while the male tenth tergum has its tip more or less incised in the other species of the P. costalis group. 21. Male tenth tergum, posterolateral corner: (0) Not produced; (1) Produced. The posterolateral corner of the male tenth tergum is more or less produced outward in most species of the P. costalis group, while it is not produced in the outgroups, P. basimaculatus, P. niger, P. sinensis, and P. stigmosus (Figs. 46, 96, 116, 124). 22. Male tenth tergum, posterolateral corner: (0) Not produced; (1) Acutely produced; (2) Roundly produced. In P. hunanensis and P. y a ng i, the posterolateral corner of the male tenth tergum is roundly produced (Figs. 83, 136). 23. Male tenth tergum: (0) Flattened; (1) Not flattened. In the outgroups, P. basimaculatus, and P. ni g er, the male tenth tergum is somewhat flattened, while in the other species of the P. c o st al is group, the male tenth tergum is not flattened. 24. Male tenth ternum: (0) Slightly narrowed toward apex; (1) Not narrowed. In the outgroups, P. niger and P. stigmosus, the male tenth tergum is slightly narrowed toward its apex (Figs. 96, 124), while in the other species of the P. costalis group, the male tenth tergum is not narrowed or even widened toward apex. 25. Male tenth sternum with anterior margin: (0) Not elevated; (1) Elevated. The male tenth sternum with the anterior margin elevated is shared by most species of the P. co s ta li s group except P. n i ge r. 26. Male tenth sternum with posterior margin medially incised: (0) Not incised; (1) Incised. In most species of the P. c o st a li s group, the male tenth sternum is medially incised on the posterior margin, while in the outgroups the posterior incision is absent and in P. basimaculatus the posterior margin is prominent medially. 27. Male tenth sternum with lateral arms: (0) Short; (1) Long. In P. acutatus, P. basiflavus, P. costalis, and P. gutianensis, the lateral arm of the male tenth sternum is long (Figs. 28, 42, 57, 79), while in the other species of the P. costalis group, the lateral arm is rather short. 28. Male tenth sternum with posterior process: (0) Absent; (1) Small; (2) Large. The posterior process is located on the sides of the posterior incision and present in a few species of the P. c os ta l is group. In P. acutatus, P. basiflavus, P. costalis, P. disjunctus, P. li i, and P. yunnanensis, the posterior process is distinct but rather small (Figs. 28, 42, 57, 64, 94, 146), while in P. stigmosus the process is abnormally enlarged (Fig. 127). 29. Male tenth sternum with lateral lobes: (0) Tuberculiform; (1) Digitiform. 30. Male tenth sternum with lateral lobes: (0) Short; (1) Long. In outgroups, P. niger, and P. stigmosus, the lateral lobes of the male tenth sternum are short (Figs. 100, 127), while they are long in the other species of the P. c os ta l is group. 31. Male tenth sternum with distal half of lateral lobes: (0) Not strongly narrowed; (1) Strongly narrowed. 32. Female eighth sternum: (0) Not narrowing toward apex; (1) Narrowing toward apex. In all the species of the P. c os t al is group, the female eighth sternum is narrowing toward apex. 33. Female eighth sternum, posterior margin: (0) Truncate; (1) Incised, forming pair of processes. In the outgroups and in the group members P. arunachalensis Ghosh, P. basimaculatus, P. fujianensis, P. gutianensis, P. hunanensis, P. triangulatus, and P. yangi, the female eighth sternum has a truncate posterior margin (Figs. 37, 52, 74, 81, 89, 134, 141), while it is incised in P. acutatus, P. c os t al is , P. disjunctus, P. orientalis, P. similis, P. s in e ns is , and P. yunnanensis. 34. Female ninth segment, sac-like lobes: (0) Absent; (1) Present. The sac-like lobe is laterally inflated from the female ninth abdominal segment, which is present in most species of the P. co s ta li s group. However, the sac-like lobe is absent in the outgroups and in P. basimaculatus. During this study, we found some females of P. arunachalensis and P. gutianensis retained an incomplete spermatphore attached to its genitalia, held pri-

6 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. marily by the lateral sac-like lobes of the ninth sternum. So, perhaps the sac-like lobes play a more important role in mating than the other female genitalic structures, and hence are much more modified and may be used as an informative character in this analysis. 35. Female ninth segment, sac-like lobes: (0) Small; (1) Large. In P. arunachalensis, P. disjunctus, P. fujianensis, P. gutianensis, P. hunanensis, P. orientalis, P. s im i li s, P. sinensis, P. yangi, and P. triangulatus, the lateral sac-like lobe of the female ninth segment is feebly developed, with its diameter about 0.5–0.75 mm at maximum (Figs. 36, 65, 73, 80, 88, 106, 113, 133, 140). In P. acutatus, P. basiflavus, P. c o s ta li s, P. similis, and P. yunnanensis, the lateral sac-like lobe is strongly developed, with its diameter up to 1.5 mm at maximum (Figs. 29, 43, 58, 147).

Results

Both of the present heuristic and branch & bound analyses resulted in only one identical most parsimonious (MP) tree (length = 67, consistency index (CI) = 0.5821, retention index (RI) = 0.7358) (Fig. 1). Its unambiguous characters have been mapped directly on the MP tree. An additional cladogram with >50% bootstrap values is shown as Figure 2. According to the MP tree and the tree with the bootstrap values, the P. co s ta li s group was found to be monophyletic. Within the ingroup, P. niger was assigned as the sister taxon with all the other group members. Similar to P. niger, three species, P. stigmosus, P. basimacualtus, and P. s in e ns is , were placed to be the sister taxa with the remaining species below their own nodes. Except for the former four species, the remaining ingroup species formed a monophyletic subgroup and separated into two smaller groups containing P. acutatus, P. basiflavus, P. c os t al is , P. disjunctus, P. li i, P. orientalis, P. si m il is , and P. hunanensis, P. yangi, P. fujianensis, and P. gutianensis. However, this subgroup is internally paraphyletic in Figure 2.

Discussion

Monopyly of the P. c o st a li s group The monophyly of the P. c os ta l is group was supported by the dorsally invisible male ninth gonostylus (char. 14: 1), the short subcylindrical male tenth tergum (char. 16: 1), the ventral tuft of the male tenth tergum (char. 19: 1), and the posterior incision of the male tenth sternum (char. 26: 1). Excluding these synapomorphic characters, the sac-like lobe of the female ninth abdominal segment might be a synapomorphic character of the ingroup, even though it is absent in P. basimaculatus. According to our recent studies on Protohermes, many species-groups are recognized, such as the P. a ss a - mensis group (Liu, Hayashi & Yang, unpublished), the P. changninganus group (Liu & Yang, 2005), the P. davidi group (Liu & Yang, 2006b), the P. differentialis group (Liu & Yang, 2006d), and the P. xanthodes group (Liu, Hayashi & Yang, 2006). The phylogenetic relationships among these species-groups have not been worked out. However, the close relationships among the P. assamensis, the P. changninganus, the P. c o st a li s , and the P. xanthodes groups might be preliminarily inferred by the widely separated ocelli, the tuft of the male tenth tergum, both of which are shared by the P. c o st a li s and P. xanthodes groups, and by the dorsally invisible male ninth gonostylus, which is shared by the P. assamensis, P. changninganus, and P. costalis groups.

Interspecific relationships of the P. costalis group The four ingroup taxa, P. n i ge r, P. stigmosus, P. basimacualtus, and P. sinensis, whose phylogenetic status is more solid inferred from the relatively high Bremer’s decay values, are generally quite distinct from each other, which implies that these species might be representatives of the remnants from former diverse lineages or have evolved for a long time.

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 7 FIGURE 1. The most parsimonious tree of the P. costalis group with the Bremer’s decay indices indicated at nodes (see text for further information). 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.

Excluding the above four ingroup taxa, the monophyletic group comprising of the other ingroups was supported by the posterior processes of the male ninth sternum close to each other (char. 12: 1) and acutely produced posterolateral corner of the male tenth tergum (char. 21: 1, char. 22: 1) of the male ninth sternum. The subclade, which contains P. si m il is , P. orientalis, P. disjunctus, P. l ii, P. yunnanensis, P. basiflavus, P. co s - talis, and P. acutatus, was supported by the posteriorly incised female eighth sternum (char. 33: 1, CI = 0.500). Within the subclade, the grouping of P. disjunctus, P. lii, P. yunnanensis, P. basiflavus, P. c o st al is , and P. acutatus was supported by the immaculate vertex (char. 3: 1, CI = 0.333) and the presence of the posterior process on the male tenth sternum (char. 28: 1, CI = 1.000). So, in P. acutatus, the weak vertex marking is considered to reflect an intermedial state during degenerating of the markings. Another subclade comprising of P. gutianensis, P. fujianensis, P. hunanensis, and P. yangi was supported by the prothorax with only one blackish vitta on each side of the pronotum (char. 4: 1, CI = 0.667). However, we consider that the relationships between P. gutianensis and the latter three species need further corrobora- tion, because several features in P. gutianensis distinctly differ from P. fujianensis, P. hunanensis, and P. yangi, but resemble P. s im i li s and P. c o st al is , such as the round lateral margins of the vertex, the indistinct wing marking patterns, and the shape of the male tenth tergum and sternum. The grouping of P. fujianensis, P. hunanensis, and P. yangi was well supported by the enlarged apical round markings on the forewings (char. 5: 1) and the strongly curved male ninth gonostylus (char. 15: 1). It is interesting that the wings are often abnormally darkened in many individuals of P. fujianensis, P. hunanensis, and P. yangi, which may also indicate a

8 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. common evolutionary history for these species, even though this character was not used in the cladistic analysis. However, considering the subgroup including 14 group members, the Bremer’s decay values of several branches are relatively low and the internal topology is paraphyletic after bootstrapping, which might be explained by these species representing a younger clade comprised of rather closely related and recently orig- inated species. Alternatively, this result might be due to the weak dispersal ability and the similarity of aquatic habitats of these corydalid species, perhaps causing a conservative morphology at the species level. Thus, the conservative morphological structures, particularly in their genitalia, might lead to considerable difficulties in finding enough informative characters for cladistic analysis.

FIGURE 2. Bootstrap 50% majority rule consensus tree (n = 1000 replicates).

Biogeography All species of the P. co s ta l is group are distributed in the Oriental Realm, except P. acutatus, which also extends to the Palaearctic Realm. Fourteen species are endemic to China, while P. arunachalensis, P. basiflavus, and P. co s ta li s are also distributed in the neighboring countries and P. disjunctus only occurs on Ishigaki and Iriomote Islands of southern Ryukyus, Japan. Considering the Chinese fauna, the species of the P. costalis group are distributed in four zoogeographical regions (Zhang, 1998), showing different interspecific distribution patterns. In the South China Region, 13 species reside (P. arunachalensis, P. basiflavus, P. basimaculatus, P. co s ta l is, P. gutianensis, P. hunanensis, P. li i , P. niger, P. orientalis, P. st ig m o s us , P. triangulatus, P. yangi, and P. yunnanensis), and nine of them (P. arunachalensis, P. basiflavus, P. basimaculatus, P. l ii , P. niger, P. stigmosus, P. triangulatus, P. yangi and P. yunnanensis) are endemic. In the Southwest China Region, there are three species, P. gutianensis, P. s i m il is , and P. yunnanensis, with P. s im i li s being endemic. The Central

FIGURES 3–5. Maps showing the geographical distributions of the species in the P. costalis group. 3. P. acutatus sp. nov. (); P. basiflavus Yang (F); P. c os tal is (Walker) (); P. disjunctus sp. nov. (9); P. lii sp. nov. (); P. yunnanensis Yang & Yang (). 4. P. arunachalensis Ghosh (); P. gutianensis Yang & Yang (); P. orientalis sp. nov. (F); P. sim il is Yang & Yang (); P. sinensis Yang & Yang (); P. stigmosus sp. nov. (); P. triangulatus sp. nov. (9). 5. P. hunanensis Yang & Yang (); P. fujianensis Yang & Yang (); P. basimaculatus sp. nov. (F); P. yangi sp. nov. (); P. n ige r Yang & Yang ().

10 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. China Region, which is the largest of the four zoogeographical regions, is divided into two main subregions, the East Hilly Plain Subregion and the Western Mountain Subregion. The former subregion contains five species, P. co s ta li s, P. fujianensis, P. gutianensis, P. orientalis, and P. sinensis, with P. fujianensis and P. s in e ns is endemic to the subregion. The latter subregion also contains five species, P. acutatus, P. c o st a li s, P. gutianensis, P. hunanensis, and P. orientalis. Protohermes acutatus is the single palaearctic representive of the P. co s - talis group in the North China Region. China, particularly the South China Region, is considered to be the distribution center of the P. costalis group since it has the richest species diversity. Previous studies on the zoogeography of Megaloptera showed that the current Asian Corydalidae fauna may originate from northeastern India or southwestern China during the split of Laurasia and Gondwanaland or the connection of the India subcontinent to Eurasia (Penny, 1993; Liu et al., 2005). Furthermore, the origination of the Asian megalopteran fauna might have happened during the split of Laurasia and Gondwanaland but not the connection of the India subcontinent to Eurasia according to the study on the generic phylogeny of Chauliodinae (Liu & Yang, 2006c). However, the center of origin of the Asian Corydalidae is corroborated by several independent studies on the Asian megalopteran fauna (Liu & Yang, 2005, 2006a; Liu et al., 2005). Considering the P. co s ta li s group, P. niger, P. st ig m o s us , P. basimaculatus, and P. triangulatus, which occupy the relatively basal phylogenetic position, were distributed in Yunnan, which consists of the western part of the South China Region and the southern part of the Southwest China Region. So, the center of origin of the P. c o st al is group might also fit with that of the Asian Corydalidae mentioned previously. The earlier speciation of the P. co s ta li s group might have happened at the mountainous region of the South China Region, especially in southern Yunnan, which is adjacent to the border between the Indian subcontinent and Laurasia. The distribution track of the P. c o s ta li s group is herein recognized from the center of origin to the northern and eastern areas. This northward dispersal may have occurred through the Southwest China Region to the North China Region, with P. acutatus as the representive of the northward invasion. Eastward dispersal may have occurred through the western parts of the South and North China Region, to the eastern parts of the South and North China Region. The occurrence of P. c o st a li s in Taiwan and P. disjunctus in the southern Ryukyus imply the eastward distribution track of the P. co s ta l is group. Thus, the close relationships among the Asian mainland, Taiwan, and southern Ryukyus might be inferred from the similar fauna of the P. costalis group.

Systematics

Genus Protohermes Weele 1907

Protohermes Weele 1907: 243. Type species Hermes anticus Walker 1853: 205, original designation. Allohermes Lestage 1927: 100. Type species Protohermes davidi Weele 1909: 254, original designation.

The Protohermes costalis species-group

Diagnosis. Median ocellus apparently transverse; posterior pair of ocelli widely separated, distance between them mostly equal to that between antennal bases. Male ninth gonostylus always enveloped by ninth tergum and invisible in dorsal view. Male tenth tergum short and subcylindrical, ventrally usually produced into a tubercle bearing a small tuft; tip usually slightly incised and densely setose. Male tenth sternum arched, basally more or less elevated at middle, apical margin usually incised medially. Female eighth sternum subtrapezoidal and narrowing toward apical portion. Female ninth abdominal segment mostly inflated laterally forming a pair of sac-like lobes.

1. Costal areas of forewings only with basal 1–3 costal cellules tinged with distinct brown markings (Fig. 9) ...... P. basimaculatus sp. nov. - Costal areas of forewings entirely hyaline or with brownish stripes in most costal cellules ...... 2 2. Head with post-ocular spine (Figs. 45, 95, 123)...... 3 - Head without post-ocular spine ...... 4 3. Pronotum dark (Fig. 95) ...... P. ni g er Yang & Yang - Pronotum mostly yellow with 5 pairs of blackish markings on each side (Figs. 123)………P. stigmosus sp. nov. 4. Forewings with brownish stripes in costal cellules ...... 5 - Forewings without any dark stripes in costal cellules (Fig. 19) ...... P. s in e ns is Yang & Yang 5. Vertex without any blackish markings...... 6 - Vertex with blackish markings near lateral sides...... 10 6. Male ninth sternum with posterior incision ...... 7 - Male ninth sternum with posterior margin truncate (Figs. 62, 144)...... 9 7. Male ninth sternum with rather shallow posterior incision (Fig. 40) ...... P. basiflavus Yang - Male ninth sternum with deep posterior incision (Figs. 55, 92)...... 8 8. Markings on pronotum large and close to each other (Fig. 53); male ninth sternum with straight lateral margin (Fig. 55) ...... P. costalis (Walker) - Markings on pronotum small and widely separated (Fig. 90); male ninth sternum with lateral margin basally arched (Fig. 92) ...... P. lii sp. nov. 9. Male ninth gonostylus feebly curved dorsad (Fig. 145); male tenth tergum with well developed ventral turbercle; female sac-like lobes on ninth abdominal segment large subquadrate (Fig. 147...... P. yunnanensis Yang & Yang - Male ninth gonostylus strongly curved dorsad (Fig. 63); male tenth tergum with feebly developed ventral turbercle; female sac-like lobes on ninth abdominal segment small suboval (Fig. 65)...... P. disjunctus sp. nov. 10. Pronotum with two pairs of blackish markings on each side ...... 11 - Pronotum with one pair of blackish vittae on each side ...... 16 11. Male ninth sternum with deep posterior incision ...... 12 - Male ninth sternum with shallow posterior incision (Fig. 103)...... P. orientalis sp. nov. 12. Male ninth sternum with V-shaped posterior incision (Fig. 130) ...... P. triangulatus sp. nov. - Male ninth sternum with trapezoidal posterior incision ...... 13 13. Forewing without yellowish round marking on apical 1/3 (Fig. 6)...... P. acutatus sp. nov. - Forewing with distinct yellowish round markings on apical 1/3...... 14 14. Forewing with median yellowish markings mostly connected (Fig. 7); male tenth tergum with posterolateral corner distinctly produced (Fig. 32) ...... P. arunachalensis Ghosh - Forewing with median yellowish markings separated (Fig. 18); male tenth tergum with posterolateral corner slightly produced (Fig. 109) ...... P. similis Yang & Yang 15. Forewings without yellowish round marking on apical 1/3 (Fig. 13) ...... P. gutianensis Yang & Yang - Forewings with distinct enlarged whitish round marking on apical 1/3 (Figs. 12, 14, 22) ...... 16 16. Legs pale yellow to brown; male ninth sternum with V-shaped posterior incision (Fig. 137)...... P. yangi sp. nov. - Legs dark; male ninth sternum with trapezoidal posterior incision...... 17 17. Mandible and antenna entirely black; lateral lobes of male tenth sternum slightly incurved (Fig. 87) ...... P. hunanensis Yang & Yang

12 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. - Mandible yellow with apical half black; antenna black with scape and pedicel yellow; lateral lobes of male tenth sternum with distal half apparently incurved (Fig. 72)...... P. fujianensis Yang & Yang

Protohermes acutatus sp. nov. (Figs. 6, 24–30)

Diagnosis. Head with two brownish or blackish markings on each side of vertex; pronotum with two blackish markings on each side; male tenth sternum with well developed V-shaped posterior incision, leaving two acutely angled, triangular distal processes. Body length 30–32 mm (male) and 43–45 mm (female); forewing length 41–42 mm (male) and 48–50 mm (female), hindwing length 37–38 mm (male) and 43–45 mm (female). Head (Fig. 24) yellowish brown, without post-ocular spine; vertex with two brownish or blackish markings on each side, lateral marking long, cuneiform, medial marking punctuate, sometimes markings indistinct. Compound eyes brown; ocelli yellowish brown, darkly margined medially. Antennae black, with scapes and pedicel yellowish brown. Mouthparts yellowish brown; maxillary palpus and labial palpus dark brown, mandible with distal half black. Prothorax (Fig. 24) yellowish brown; pronotum with two blackish markings on each side. Meso- and metathorax pale yellow to yellowish brown, each dorsally with one pair of brownish markings laterally. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs yellow, with dense, yellowish, short setae; tibiae black apically; tarsal claws reddish brown. Forewings (Fig. 6) pale brown, with 1 large yellowish marking at base, 3–4 yellowish markings at middle, and 1 extremely small white spot on apical 1/3; costal cellules with indistinct brownish stripes. Hindwings paler than forewings, with 1 extremely small white spot on apical 1/3. Veins dark, except veins in yellowish markings pale yellow. Rs 9 to10-branched, last branch trifurcate; 9–10 crossveins between R1 and Rs; M1+2 6 to7-branched, M3+4 2 to 4-branched; 1A 3-branched. Abdomen brown with yellowish short setae. Male ninth tergum (Fig. 25) subquadrate, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 26) broad; posterior margin with deep trapezoidal incision, forming two acutely angled, triangular distal processes; central portion inflated. Ninth gonostylus (Fig. 27) unguiform and slightly curved inwards and dorsad. Tenth tergum (Figs. 25–26) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner slightly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 28) arched, anteriorly distinctly elevated at middle, posterior incision well developed and V-shaped, forming two acutely angled triangular processes; lateral lobes elongated and digitiform, distally slightly incurved. Female eighth sternum (Fig. 29–30) strongly sclerotized and subtriangular in lateral view; apical margin distinctly elevated with arcuate median incision, forming pair of obtuse processes; ninth segment (Fig. 29) laterally inflated in one pair of large subquadrate sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one round lower lobe. Type materials. Holotype %, CHINA: Chongqing, Wanxian, Wangerbao, 1200 m, 27.V.1994, Wenzhu Li (IZCAS). Paratypes 3 % &, CHINA: Chongqing, Wanxian, Wangerbao, 1200 m, 28.V.1994, Wenzhu Li (IZCAS); %, CHINA: Chongqing, Changshou, Nanmuyuan, 450 m, 9.VI.1994, Youwei Zhang (IZCAS); %, CHINA: Chongqing, Changshou, Nanmuyuan, 450 m, 9.VI.1994, Wenzhu Li (IZCAS); 2 &, CHINA: Chong- qing, Fengdu, Shiping, 610 m, 2.VI.1994, Youwei Zhang (IZCAS); &, CHINA: Chongqing, Fengdu, Shiping, 610 m, 3.VI.1994, Youwei Zhang (IZCAS); %, CHINA: Shannxi, Langao, Minzhu, 4.VII.2003, Caixia Yuan & Yushuang Liu (HBU). Distribution. China (Chongqing, Shannxi). Etymology. The specific epithet ‘acutatus’ refers to the acutely angled apical processes of the male tenth sternum.

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 13 FIGURES 6–15. Habitus of the species in the P. costalis group. 6. P. acutatus sp. nov. (Holotype), male. 7. P. arunachalensis Ghosh, male. 8. P. basiflavus Yang (Holotype), male. 9. P. basimaculatus sp. nov. (Holotype), male. 10. P. c os tal is (Walker), male. 11. P. disjunctus sp. nov. (Holotype), male. 12. P. fujianensis Yang & Yang (Holotype), male. 13. P. gutianensis Yang & Yang (Holotype), male. 14. P. hunanensis Yang & Yang (Holotype), male. 15. P. lii sp. nov. (Holotype), male. Scale bars = 5 mm.

14 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. FIGURES 16–23. Habitus of the species in the P. costalis group. 16. P. niger Yang & Yang (Holotype), male. 17. P. o ri- entalis sp. nov. (Holotype), male. 18. P. similis Yang & Yang (Holotype), male. 19. P. si ne n sis Yang & Yang (Holotype), male. 20. P. stigmosus sp. nov. (Holotype), male. 21. P. triangulatus sp. nov. (Holotype), male. 22. P. y an gi sp. nov. (Holotype), male. 23. P. yunnanensis Yang & Yang (Holotype), male. Scale bars = 5 mm.

Remarks. The new species appears to be closely related to P. s im il is in having a similar pattern of markings on the head and prothorax, while it is distinguished by the pattern of markings on the wings and the structure of the male genitalia. In P. acutatus, the yellowish round markings on the apical 1/3 of the forewings are absent, instead having an extremely small white spot, and the posterior incision of the male tenth sternum is distinct, forming a pair of acutely angled posterior processes. In P. s im i li s, the round yellowish markings are distinct on the apical 1/3 of the forewings, and the posterior incision of the male tenth sternum is extremely small.

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 15 FIGURES 24–30. Protohermes acutatus, sp. nov. 24. Male head and pronotum, dorsal view. 25. Male genitalia, dorsal view. 26. Male genitalia, ventral view. 27. Male ninth gonostylus, caudal view. 28. Male tenth sternum, ventral view. 29. Female genitalia, lateral view. 30. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes arunachalensis Ghosh (Figs. 7, 31–37)

Protohermes arunachalensis Ghosh, 1991. Rec. Zool. Surv. India, 88: 147. Type locality: India (Arunachal Pradesh).

Diagnosis. Head with 2–4 pairs of black markings; pronotum with two pairs of large black markings; male ninth sternum with trapezoidal posterior incision; tenth tergum with outer-distal corner produced. Body length 29–35 mm (male) and 43–47 mm (female); forewing length 40–45 mm (male) and 48–50 mm (female), hindwing length 34–40 mm (male) and 42–45 mm (female). Head (Fig. 31) yellow, without post-ocular spine; vertex with 2–4 pairs of black markings, posterolateral markings subtriangular. Occiput laterally with pair of black markings. Compound eyes greyish brown; ocelli yellow, darkly margined medially. Antennae dark, with scapes and pedicel yellow. Mouthparts yellow; mandible with distal half black. Thorax yellow. Pronotum (Fig. 31) laterally with two pairs of large black markings; posterolateral corner black. Meso- and metanota brownish on lateral sides. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs yellow with dense yellowish short setae; fore and middle tibiae with dorsal surface black,

16 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. hind tibia occasionally with dorsal surface also black; tarsi dark; tarsal claws reddish brown. Forewings (Fig. 7) pale brown, with 1 subtriangular, large yellowish marking at base, several fused yellowish markings at middle, and 1 round yellowish marking on apical 1/3; costal cellules with distinct brown stripes. Hindwings much paler than forewings, with basal half almost hyaline, and with 1 round yellowish marking on apical 1/3. Veins pale yellow but dark toward apex. Rs 10-branched, last branch bifurcate; 7–12 crossveins between R1 and Rs;

M1+2 6 to 9-branched, M3+4 3 to 4-branched; 1A 3-branched.

FIGURES 31–37. Protohermes arunachalensis Ghosh. 31. Male head and pronotum, dorsal view. 32. Male genitalia, dorsal view. 33. Male genitalia, ventral view. 34. Male ninth gonostylus, caudal view. 35. Male tenth sternum, ventral view. 36. Female genitalia, lateral view. 37. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen yellowish brown with yellowish short setae. Male ninth tergum (Fig. 32) subquadrate, with arched anterior and slightly incised posterior margin. Ninth sternum (Fig. 33) broad with arched lateral margin, posterior margin with trapezoidal incision forming two blunt triangular posterior processes; central portion apparently inflated. Ninth gonostylus (Fig. 34) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 32–33) short subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 35) arched, anteriorly slightly elevated at middle; posterior margin medially with one extremely small incision; lateral lobes thick and long digitiform. Female eighth sternum (Figs. 36–37) strongly sclerotized, subtriangular in lateral view, posterior margin truncate; ninth segment (Fig. 36) laterally with a pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one short subtriangular upper and one semicircular lower lobe.

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 17 Materials examined. 4 % 4 &, CHINA: Yunnan, Baoshan, Baihualing, 1500 m, 23.V.2006, Xingyue Liu (CAU); 6 % 2 &, CHINA: Yunnan, Baoshan, Baihualing, 1500 m, 22.V.2006, Xingyue Liu (CAU); 2 % &, CHINA: Yunnan, Tengchong, Longchuanjiang, 1050 m, 16.V.2006, Xingyue Liu (CAU); 3 %, CHINA: Yun- nan, Tengchong, Jietou, 1500 m, 17.V.2006, Xingyue Liu (CAU). Distribution. China (Yunnan); India (Arunachal Pradesh). Remarks. Protohermes arunachalensis was originally described by Ghosh (1991) based on a single male from Arunachal Pradesh, India. Ghosh (1991) also noted that this species closely resembles Protohermes anticus (Walker). Although the type specimen was not available for examination, the presence of a subcylindrical male tenth tergum with a ventral tuft according to the original illustration easily places this species in the P. costalis group. During our survey of western Yunnan, China, several individuals were found and identified as P. arunachalensis, agreeing closely with the original description. The redescription of the male and description of the female was based on the specimens from western Yunnan. This species seems to be closely related to P. triangulatus, but can be separated from it by the trapezoidal posterior incision of the male ninth sternum. In P. triangualtus, the male ninth sternum bears a V-shaped posterior incision. In the original description of P. arunachalensis, a black squarish marking at the tip of the hindwing was mentioned, which is absent from our specimens as well as in the other species of the P. co s ta li s group. However, whether the black marking is a sta- ble character or resulted from an aberrant individual should be investigated.

Protohermes basiflavus Yang (Figs. 8, 38–44)

Protohermes basiflavus Yang et al., 2004. Insects from Mt. Shiwandashan Area of Guangxi, p. 265. Type locality: Guangxi (Fangcheng, Napo).

Diagnosis. Head without any dark markings; pronotum with two large blackish vittae on each side; male ninth sternum with rather shallow trapezoidal posterior incision. Body length 28–34 mm (male) and 45 mm (female); forewing length 45–48 mm (male) and 55 mm (female), hindwing length 40–43 mm (male) and 52 mm (female). Head (Fig. 38) yellowish brown, without post-ocular spine; vertex without markings. Compound eyes pale brown; ocelli yellow, darkly margined medially. Antennae dark, with yellow scapes and brown pedicel. Mouthparts yellow; mandible with distal half dark. Prothorax yellow; pronotum (Fig. 38) with two pairs of large blackish markings on each side. Meso- and metathorax yellowish brown, each with one pair of brownish markings laterally. Thoracic pilosity yellow, longer on meso- and metathorax. Legs yellow with dense, yellowish, short setae; tibiae and tarsi black, with narrow bases of fore and middle tarsi and basal 1/3 of hind tarsi yellowish brown; tarsal claws reddish brown. Forewings (Fig. 8) pale smoky brown, with 1 large yellowish marking at base, 3–4 mostly connected yellowish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with distinct brown stripes. Hindwings much paler than forewings, respectively with 1 round yellowish marking at middle and apical 1/3. Veins dark except veins in yellowish markings yellow. Rs 10-branched, last branch trifurcate; 10–

11 crossveins between R1 and Rs; M1+2 6-branched, M3+4 4-branched; 1A 3-branched. Abdomen pale brown with yellowish short setae. Male ninth tergum (Fig. 39) subquadrate, with trapezoidal anterior margin and truncate or sightly incised posterior margin. Ninth sternum (Fig. 40) broad with arched lateral margin, posterior margin with rather shallow trapezoidal incision, forming two blunt posterior processes; central portion apparently inflated. Ninth gonostylus (Fig. 41) unguiform and slightly curved dorsomedially, sometimes with an additional dent near tip. Tenth tergum (Figs. 39–40) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 42) arched, anteriorly slightly

18 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. elevated at middle; posterior incision extremely small and V-shaped, forming two papillated processes; lateral lobes digitiform and slightly incurved. Female eighth sternum (Figs. 43–44) strongly sclerotized, subtriangular in lateral view, posterior margin distinctly elevated with distinct median incision, forming pair of obtuse processes; ninth segment (Fig. 43) laterally inflated in one pair of large subquadrate sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one semicircular lower lobe. Material examined. Holotype %, CHINA: Guangxi, Fangcheng, Fulong, 500 m, 24–25.V.1999, Dajun Liu & Xuezhong Zhang (IZCAS). Paratypes: &, CHINA: same data as holotype (IZCAS); %, CHINA: Guangxi, Napo, Defu, 1350 m, 18.VI.2000, Jun Chen (IZCAS); %, CHINA: Guangxi, Napo, Beidou, 500 m, 22.VI.2000, Jun Chen (IZCAS). %, CHINA: Guangxi, Shiwandashan, 19.V. 2006, Kuiyan Zhang (CAU); %, CHINA: Yunnan, Funing, 800 m, 18.V.1979, Yaoxuan Yin (CAU); %, BURMA: Dawna, Sud-qst, 25.XI.1997, Fumio Hayashi (CAU). Distribution. Burma (Dawna); China (Guangxi, Yunnan). Remarks. This species appears to be closely related to P. costalis in having similar immaculate head and wing patterns, but can be easily separated by the rather shallow posterior incision of the male ninth sternum. In P. costalis, the posterior incision of the male ninth sternum is deep.

FIGURES 38–44. Protohermes basiflavus Yang. 38. Male head and pronotum, dorsal view. 39. Male genitalia, dorsal view. 40. Male genitalia, ventral view. 41. Male ninth gonostylus, caudal view. 42. Male tenth sternum, ventral view. 43. Female genitalia, lateral view. 44. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes basimaculatus sp. nov. (Figs. 9, 45–52)

Diagnosis. Head with two pairs of black markings; pronotum with two pairs of small, widely separated black

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 19 markings; costal areas of forewing with basal 1–3 cellules marked brown; male tenth tergum laterally flattened; female ninth abdominal segment without sac-like lobes. Body length 25–32 mm (male) and 35 mm (female); forewing length 37–40 mm (male) and 50 mm (female), hindwing length 34–35 mm (male) and 44 mm (female). Head (Fig. 45) yellow, with blunt post-ocular spine; vertex with two pairs of black markings, lateral markings subtriangular, medial markings suboval. Occiput laterally with pair of black markings. Compound eyes greyish brown; ocelli yellow, darkly margined medially. Antennae dark, with scapes and pedicel yellow. Mouthparts yellow; mandible with distal half black.

FIGURES 45–52. Protohermes basimaculatus, sp. nov. 45. Male head and pronotum, dorsal view. 46. Male genitalia, dorsal view. 47. Male genitalia, ventral view. 48. Male genitalia, lateral view. 49. Male ninth gonostylus, caudal view. 50. Male tenth sternum, ventral view. 51. Female genitalia, lateral view. 52. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

20 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. Thorax yellow. Pronotum (Fig. 45) laterally with two pairs of small, widely separated black markings, posterior marking much broader than anterior marking, but sometimes separated into two narrow markings; posterolateral corner black. Meso- and metanota darker on lateral sides, medially pale. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs yellow with dense yellowish short setae; 5th tarsus dark; tarsal claws reddish brown. Forewings (Fig. 9) slightly smoky brown, with 1 large yellowish marking at base, several fused yellowish markings and some small markings on crossveins between R1 and Rs at middle, and 1 round yellowish marking on apical 1/3; costal areas hyaline except for basal 1–3 cellules with distinct brown markings. Hindwings much paler than forewings, with 1 rather indistinct round yellowish marking on apical 1/3. Veins pale yellow but slightly darker toward apex; costal crossveins dark. Rs 9 to 10-branched, last branch bifurcate; 9–11 crossveins between R1 and Rs; M1+2 5-branched, M3+4 2-branched; 1A 3-branched. Abdomen yellow with yellowish short setae. Male ninth tergum (Fig. 46) subquadrate, with arched anterior and slightly incised posterior margin. Ninth sternum (Fig. 47) broad with nearly straight lateral margin, posterior margin with wide V-shaped incision, forming two acutely angled triangular posterior processes; central portion apparently inflated. Ninth gonostylus (Fig. 48) unguiform and strongly curved dorsomedially. Tenth tergum (Figs. 46–48) short subcylindrical, laterally flattened, ventrally with one tuft at posteromedial corner; tip slightly incised and densely setose. Tenth sternum (Fig. 50) arched, anteriorly slightly elevated at middle; posterior margin medially produced with one small incision at tip; lateral lobes long digitiform. Female eighth sternum (Figs. 51–52) strongly sclerotized, subtriangular in lateral view, posterior margin truncate; ninth segment (Fig. 51) without sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one short digitiform upper and one semicircular lower lobe. Type materials. Holotype %, CHINA: Yunnan, Baoshan, Baihualing, 1500 m, 22.V.2006, Xingyue Liu (CAU). Paratypes: 2 %, CHINA: same data as holotype (CAU); 2 % &, CHINA: Yunnan, Baoshan, Baihual- ing, 1500 m, 23.V.2006, Xingyue Liu (CAU). Distribution. China (Yunnan). Etymology. The specific epithet ‘basimaculatus’ refers to the feature of the forewing with only the basal 1–3 costal cellules marked brown in the new species. Remarks. The new species is distinguished from the other species of the P. costalis group by the forewing only with the basal 1–3 costal cellules marked brown, the laterally flattened male tenth tergum, and the female ninth abdominal segment without sac-like lobes.

Protohermes costalis (Walker) (Figs. 10, 53–59)

Hermes costalis Walker, 1853. Cat. Brit. Mus. Neur., p. 207. Type locality: N. China. Protohermes griseus Stitz, 1914. Berlin Sitzb. Ges. Naturf. 5: 201. Type locality: Formosa.

Diagnosis. Head without dark markings; pronotum with two blackish vittae on each side; male ninth sternum with deep trapezoidal posterior incision, leaving two acute-angled triangular posterior processes. Body length 30–34 mm (male) and 45–52 mm (female); forewing length 36–48 mm (male) and 45–51 mm (female), hindwing length 32–41 mm (male) and 42–45 mm (female). Head (Fig. 53) yellowish brown, without post-ocular spine; vertex without any markings. Compound eyes pale yellowish brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel yellow. Mouthparts yellow or yellowish brown; mandible with distal half black. Thorax yellow. Pronotum (Fig. 53) with two pairs of blackish markings on each side. Meso- and metanota each with one pair of brownish markings on lateral sides. Thoracic pilosity yellow, much longer on meso- and

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 21 meta thorax. Legs yellow with dense yellowish short setae; tibiae and tarsi mostly dark; tarsal claws reddish brown. Forewings (Fig. 10) pale smoky brown, with 1 large yellowish marking at base, 3–4 mostly connected yellowish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with distinct brown stripes. Hindwings much paler than forewings, with 1 round yellowish marking on apical 1/3. Veins yellowish brown except veins in yellowish markings yellow. Rs 10-branched, last branch bifurcate or trifurcate; 11 crossveins between R1 and Rs; M1+2 5 to 6-branched, M3+4 4-branched; 1A 3-branched. Abdomen brown with yellowish short setae. Male ninth tergum (Fig. 54) subquadrate, with arched anterior and posterior margin. Ninth sternum (Fig. 55) broad with nearly straight lateral margin, posterior margin with deep trapezoidal incision, forming two acutely angled triangular posterior processes; central portion inflated. Ninth gonostylus (Fig. 56) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 54–55) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 57) arched, anteriorly slightly elevated at middle; posterior incision extremely small and V-shaped, forming two papillated processes; lateral lobes digitiform with apical half strongly narrowing, incurved. Female eighth sternum (Figs. 58–59) strongly sclerotized, subtriangular in lateral view, posterior margin distinctly elevated with shallow median incision, forming pair of obtuse processes; ninth segment (Fig. 58) laterally inflated in one pair of large suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one subquadrate lower lobe.

FIGURES 53–59. Protohermes costalis (Walker). 53. Male head and pronotum, dorsal view. 54. Male genitalia, dorsal view. 55. Male genitalia, ventral view. 56. Male ninth gonostylus, caudal view. 57. Male tenth sternum, ventral view. 58. Female genitalia, lateral view. 59. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Material examined. CHINA: &, Yunnan, Weixin, 20.VI.1980 (CAU); &, Guizhou, 10.V.1963 (CAU); % &, Guizhou, Sinan, VI.1973, Guiqing Wang (CAU); %, Guangxi, Jinxiu, 10.VI.1982, Chikun Yang (CAU); &,

22 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. Guangxi, Lingchuan, Lingtian, 4.VI.1984, Ding Yang (CAU); 2 &, Guangxi, Jinxiu, Luoxiang, 200 m, 15.V.1999, Xuezhong Zhang (IZCAS); % &, Guangxi, Jinxiu, Luoxiang, 200 m, 15.V.1999, Hui Xiao (IZCAS); &, Guangxi, Tian-e, Buliuhe, 6.VIII.2002, Ding Yang (CAU); 3 % 6 &, Guangxi, Maoershan, 900 m, 2.VII.2003, Xingyue Liu (CAU); %, Guangdong, Dinghushan, 22.IV.1965, Youwei Zhang (IZCAS); &, Guangdong, Conghua, IV.1976, Rongzong Wu (CAU); 3 &, Guangdong, Chebaling, 19–21.IV.2002 (CAU); &, Fujian, Chong-an, Xingcun, Sangang, 740 m, 16.VII.1960, Yiran Zhang (IZCAS); %, Fujian, Shaowu, Tongmu, 28.VII.1979, Shimei Song (IZCAS); &, Fujian, Sangang, 10.VIII.1979, Rongquan Cai (IZCAS); &, Fujian, Sangang, 17.VIII.1979 (IZCAS); &, Fujian, 20.VIII.1981 (CAU); %, Fujian, Jiangle, Longqishan, 650 m, 27.VI.1991, Longlong Yang (IZCAS); 2 &, Fujian, Jiangle, Longqishan, 700 m, 1–6.VIII.1991, Shimei Song (IZCAS); %, Zhejiang, Jinhua, 21.VI.1980 (IZCAS); &, Zhejiang, Lishui, Wanxiangshan, 12.IX.1980 (CAU); 5 &, Zhejiang, Tianmushan, 21–22.VII.1973, Peiyu Yu (CAU); %, Zhejiang, Tianmushan (IZCAS); %, Zhejiang, Longwangshan, 490 m, 12–14.VI.1996, Chikun Yang (CAU); 2 % 5 &, Hunan, Nanyue, Mojingtai, 20–23.VI.1963, Chikun Yang (CAU); &, Hunan, Hengshan, 12.VII.1980 (IZCAS); 2 &, Hunan, Yongshun, Shanmuhe Forestry Center, 600 m, 4–7.VIII.1988 (IZCAS); 3& Hubei, Lichuan, 810 m, 20–23.VII.1989, Shuyong Wang (IZCAS); %, Jiangxi, Arboretum of Lushan, 8.VIII.1975 (IZCAS); 2 %, Jiangxi, Guling, 13– 17.VII.1935 (IZCAS); %, Jiangxi, Guling, 2.VIII.1935 (IZCAS); 2 %, Anhui, Huangshan, Wenquan, 17– 20.VII.1977, Fasheng Li (CAU); 4 % 3 &, Henan, Shangcheng, Huangbaishan, 200 m, 14.VII.1999, Xiaocheng Shen & Yingdang Ren (CAU); % &, Taiwan, Wulai, 5.V.1994, T. Shimizu (HC); 2 &, Taiwan, Wulai, 29.VI.1987, Fumio Hayashi (HC); %, Taiwan, Wulai, emerged from reared larva, Fumio Hayashi (CAU); &, Taiwan, emerged from reared larva in 1988, Fumio Hayashi (CAU). Distribution. China (Anhui, Fujian, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangxi, Tai- wan, Yunnan, Zhejiang); India (Assam). Remarks. After the revision by Kimmins (1948), P. co s ta li s is considered to be a valid species and not a synonym of Hermes anticus Walker. For comparison of P. costalis with other related species, see the remarks under P. yunnanensis.

Protohermes disjunctus sp. nov. (Figs. 11, 60–66)

Diagnosis. Head yellow without any markings; pronotum with two pairs of widely separated black markings; wings with several yellowish markings; male ninth sternum with posterior margin slightly incised or even truncate; male tenth tergum acutely produced on posterolateral portion, ventrally with feebly developed tufted tubercle; female abdominal segment with pair of small lateral lobes. Body length 20–28 mm (male) and 27–32 mm (female); forewing length 24–33 mm (male) and 33–41 mm (female), hindwing length 21–29 mm (male) and 30–35 mm (female). Head (Fig. 60) yellow without any markings; post-ocular spine absent. Compound eyes grayish brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel yellow. Mouthparts yellow; mandible with distal half blackish to reddish brown. Prothorax (Fig. 60) yellow; pronotum with two pairs of black markings near lateral margins, posterior pair much larger. Meso- and metathorax pale yellow. Thoracic pilosity pale yellow, much longer on meso- and metathorax. Legs pale yellow with short, dense, yellowish setae; foreleg with wide apex of tibiae dark, tarsi dark, tarsal claws reddish brown. Forewings (Fig. 11) slightly grayish brown, with 1 large subtriangular yellowish marking at base, 1 large round and several small discrete yellowish markings at middle, and 1 round yellowish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings slightly grayish brown except basal half hyaline, respectively with 1 round yellowish marking at middle and apical 1/3. Veins yellowish brown, much darker on apical half, but much paler in yellowish markings. Rs 8 to 10-branched, last

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 23 branch bifurcate or trifurcate; 8 crossveins between R1 and Rs; M1+2 6-branched, M3+4 2-branched; 1A 3- branched. Abdomen yellowish brown. Male ninth tergum (Fig. 61) subtrapezoidal, with arched anterior margin and nearly truncate posterior margin. Ninth sternum (Fig. 62) subquadrate, broader than ninth tergum; lateral margins slightly arched; posterior margin slightly incised or even truncate; central portion apparently inflated. Ninth gonostylus (Fig. 63) short, unguiform, strongly curved dorsomedially. Tenth tergum (Figs 61–62) short, subcylindrical, its posterolateral corner distinctly and acutely produced, its ventral portion with one feebly developed tufted tubercle near posteromedial corner; its tip slightly incised and densely setose. Tenth sternum (Fig. 64) arched, anterior margin slightly elevated, posterior margin produced into pair of roundly angled processes; lateral lobes somewhat thickened, digitiform, slightly incurved. Female eighth sternum (Figs. 65–66) strongly sclerotized, subtriangular in lateral view, with posterior margin distinctly incised, forming pair of roundly produced processes; ninth segment laterally inflated in pair of small suboval sac-like lobes (Fig. 65); ninth gonocoxite broad, feebly pointed posteriorly, with small digitiform processes at tip; tenth tergum short, with posterior margin medially incised, leaving one thick digitiform dorsal lobe and one semicircular ventral lobe.

FIGURES 60–66. Protohermes disjunctus, sp. nov. 60. Male head and pronotum, dorsal view. 61. Male genitalia, dorsal view. 62. Male genitalia, ventral view. 63. Male ninth gonostylus, caudal view. 64. Male tenth sternum, ventral view. 65. Female genitalia, lateral view. 66. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Type materials. Holotype %, JAPAN: Iriomote Island, Urauchi-gawa, 23.V.1995, Shin-ichi Suda (CAU). Paratypes: %, the same data as holotype (HC); &, JAPAN: Iriomote Island, Takana-gawa, 25.IV.1988, Fumio Hayashi (CAU). Others material: JAPAN: 5 % 2 &, Iriomote Island, Takana-gawa, emerged from reared larvae in 1987, 1988, Fumio Hayashi (HC); 4 % 3 &, Ishigaki Island, Nagura-gawa, emerged from reared larvae in 1988, Fumio Hayashi (HC).

24 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. Distribution. Japan (Ishigaki Island and Iriomote Island). Etymology. The specific epithet ‘disjunctus’ refers to the geographic distribution of the new species, which is endemic to the Ishigaki and Iriomote Islands of Japan. Remarks. Hayashi (1989b) assumed that this species (his Protohermes sp.) might be divided from P. co s - talis during the separation of the Iriomote Island from Taiwan. However, considering the genitalia, P. disjunctus appears to be closely related to P. yunnanensis in having a similar male ninth sternum, but it can be easily separated from P. yunnanensis by the male tenth tergum with the feebly developed tufted tubercle and the small female sac-like lobes. In P. yunnanensis, the male tenth tergum is ventrally produced into one strongly developed tufted tubercle, and the female sac-like lobes are rather large.

Protohermes fujianensis Yang & Yang (Figs. 12, 67–74)

Protohermes fujianensis Yang & Yang, 1999. Fauna of Insects in Fujian Province of China, Vol. 3, p. 171. Type locality: Fujian (Chong-an).

Diagnosis. Head with one pair of brown markings laterally; pronotum with one pair of dark vittae on lateral margins; forewings pale brown with several whitish markings; male ninth sternum with trapezoidal posterior incision; male tenth sternum with lateral lobes distinctly incurved. Body length 27–29 mm (male) and 30–31 mm (female); forewing length 36–37 mm (male) and 41–45 mm (female), hindwing length 32–34 mm (male) and 36–41 mm (female). Head (Fig. 67) yellowish brown without post-ocular spine, dorsally with one pair of brown markings on lateral sides. Compound eyes brown; ocelli pale yellow, darkly margined medially. Antennae black with scapes and pedicel pale yellow. Mouthparts yellow; mandible with distal half black. Occiput yellowish brown with one pair of brownish markings laterally. Prothorax (Fig. 67) pale yellow; pronotum with one pair of dark vittae on lateral margins, sometimes nar- rowly divided midway. Meso- and metathorax yellow. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs dark with dense yellowish short setae; coxae, trochanters and basal tibiae pale yellow; tarsal claws reddish brown. Forewings (Fig. 12) pale brown, with 1 round whitish marking at base, 2 round whitish markings at middle and 1 distinctly enlarged, whitish round marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings pale brown except basal half subhyaline, respectively with 1 round whitish marking at base, middle and apical 1/3. Veins pale brown, except veins in whitish markings pale yellow. Rs 9 to 11-branched, last branch bifurcate; 10 crossveins between R1 and Rs; M1+2 4 to 8-branched, M3+4 2 to 4- branched; 1A 3-branched. Abdomen dark with yellowish short setae. Male ninth tergum (Fig. 68) subtrapezoidal, with arched anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 69) broad with arched lateral margin, posterior margin with shallow trapezoidal incision, forming two large roundly produced apical processes; central portion inflated. Ninth gonostylus (Fig. 70) unguiform, curved dorsomedially. Tenth tergum (Fig. 71) short and looks more trapezoidal, ventrally with one feebly developed, tufted tubercle at middle; posterolateral corner distinctly and acutely produced, tip slightly incised and densely setose. Tenth sternum (Fig. 72) arched, anteriorly with one subtriangular process at middle; posterior incision indistinct; lateral lobes digitiform with distal half incurved. Female eighth sternum (Figs. 73–74) strongly sclerotized and subtriangular in lateral view, with truncate posterior margin; ninth segment (Fig. 73) laterally inflated in one pair of small oval sac-like lobes; ninth gonocoxite valvate and membranous with a digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one round lower lobe. Material examined. Holotype %, CHINA: Fujian, Chong-an, 1.VII.1980 (CAU). %, CHINA: Fujian,

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 25 10.V.1982 (CAU); %, CHINA: Fujian, 3.VI.1989 (CAU); &, CHINA: Fujian, 5.V.1988, Liying Zhao (CAU); &, CHINA: Fujian, 1.VI.1989 (CAU); % &, CHINA: Fujian, no detailed data (CAU). Distribution. China (Fujian). Remarks. Protohermes fujianensis appears to be closely related to P. hunanensis and can be distinguished from the latter as follows: mandible yellow with distal half black, antenna black with scape and pedicel yellow, male tenth sternum with lateral lobes incurved in apical half. By contrast, P. hunanensis has the mandible and antenna entirely black and feebly incurved lateral lobes of the male tenth sternum.

FIGURES 67–74. Protohermes fujianensis Yang & Yang. 67. Male head and pronotum, dorsal view. 68. Male genitalia, dorsal view. 69. Male ninth sternum, ventral view. 70. Male ninth gonostylus, caudal view. 71. Male tenth tergum, ventral view. 72. Male tenth sternum, ventral view. 73. Female genitalia, lateral view. 74. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes gutianensis Yang & Yang (Figs. 13, 75–81)

Protohermes gutianensis Yang & Yang, 1995b. Insects and macrofungi of Gutianshan, Zhejiang, p. 129. Type locality: Zhejiang (Kaihua).

Diagnosis. Head with black markings from anterior edge of compound eyes through to occiput and a narrow blackish vittae medially on vertex; pronotum with one pair of blackish vittae on lateral margins; wings pale

26 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. smoky brown with several indistinct yellowish spots except for apical regions without spots; male ninth sternum with suboblong posterior incision. Body length 28–31 mm (male) and 40–45 mm (female); forewing length 38–48 mm (male) and 48–52 mm (female), hindwing length 35–41 mm (male) and 43–47 mm (female). Head (Fig. 75) yellowish brown, without post-ocular spine, dorsally black near post-ocular margins, sometimes entirely black near lateral margins; vertex also with one narrow blackish vittae medially. Com- pound eyes brown; ocelli yellow, darkly margined medially. Antennae black with scapes and pedicel brown. Mouthparts yellow; mandible with distal half black. Prothorax (Fig. 75) yellowish brown; pronotum with one pair of blackish vittae laterally. Meso- and metathorax yellow, each dorsally with one pair of brownish spots on anterolateral corners. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs black with dense, pale yellow, short setae; coxae and trochanters pale yellow; femora brown; tarsal claws reddish brown. Forewings (Fig. 13) slightly smoky brown, with 1 small yellowish marking at base, 2–4 yellowish markings along cubital veins at middle; costal cellules with distinct brownish stripes. Hindwings much paler than forewings with hyaline basal half. Veins pale brown, except pale yellow veins in yellowish markings and base of hindwings. Rs 9–10 branched, last branch mostly trifurcate; 9 to 10-crossveins between R1 and Rs; M1+2 5 to 7-branched, M3+4 3 to 4-branched; 1A 3-branched. Abdomen brown with short, yellow setae. Male ninth tergum (Fig. 76) subquadrate, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 77) broad with arched lateral margin; posterior margin with deep suboblong incision, forming two broad acute-angled subtriangular processes; central portion inflated. Ninth gonostylus (Fig. 78) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 76–77) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 79) arched, anteriorly with one trapezoidal process at middle and posteriorly with one small triangular incision; lateral lobes slender digitiform and slightly incurved. Female eighth sternum (Figs. 80–81) strongly sclerotized, subtrapezoidal in lateral view, with truncate posterior margin; ninth segment (Fig. 80) laterally inflated in one pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one long digitiform upper and one short round lower lobe. Material examined. Holotype %, CHINA: Zhejiang, Gutianshan, 28.VII.1992, Hong Wu (CAU). CHINA: &, Zhejiang, Lishui, Dashanfeng, 3.VII.1979 (CAU); 2 %, Zhejiang, Longwangshan, 490 m, 12–14.VI.1996, Chikun Yang (CAU); %, Zhejiang, Tianmushan, 500 m, 26.VII.2000, Siqin Ge (IZCAS); &, Guangxi, Hua- ping, Cujiang, Chikun Yang (CAU); 2 % &, Guangxi, Jinxiu, 720 m, 11.VI.1983, Xinli Wang (CAU); %, Guangxi, Jinxiu, 720 m, 6.VI.1983, Fasheng Li (CAU); %, Guangxi, Jinxiu, 720 m, 10.VI.1983, Fasheng Li (CAU); 4 &, Guangxi, Guilin, Maoershan, 900 m, 2.VII.2003, Xingyue Liu (CAU); 3 &, Guangdong, Lian- ping, 16.V.1973 (IZCAS); &, Guangdong, Conghua, VI.1976, Rongzong Wu (CAU); &, Fujian, Shaowu, 13.V.1978 (CAU); %, Fujian, Chong-an, Sangang, 17.VIII.1979 (IZCAS); %, Fujian, Meihuashan, Qiushan, 24.VII.1988, Chikun Yang (CAU); 2 &, Fujian, Jiangle, Longqishan, 25.V.1991, Yongshan Shi (IZCAS); &, Fujian, Jiangle, Longqishan, 21.V.1991, Yongshan Shi (IZCAS); &, Fujian, Jiangle, Longqishan, 17.V.1991, Wenzhu Li (IZCAS); &, Guizhou, Sinan, VI.1973, Guiqing Wang (CAU); % &, Guizhou, Fanjingshan, 600 m, 3.VI.2002, Ding Yang (CAU); &, Guizhou, Jiangkou, Fanjingshan, 11.VII.1988, Xiaochun Zhang (IZCAS); %, Guizhou, Shiqian, Jinwu, 600–800 m, 22.VII.1988, Hongxing Li (IZCAS); &, Guizhou, Leishan, Taojiang, 1000 m, 5.VII.1998, Xiaochun Zhang (IZCAS); %, Sichuan, Guanxian, 700–1000 m, 27.V.1963, Xuezhong Zhang (IZCAS); &, Sichuan, Pengshui, Taiyuan, 800 m, 12.VII.1989, Longlong Yang (IZCAS); &, Hunan, Yiyang, 22.VI.1981 (IZCAS); &, Jiangxi, Yifeng, 7.VI.1959 (IZCAS); % 3 &, Gansu, Kangxian, Lianghe, 800 m, 30.VII.1980, Chunhua Yang (CAU); % 2 &, Gansu, Kangxian, Qinghelinchang, 1400 m, 7.VII.1999, Tongli He (IZCAS); 4 % 6 &, Henan, Shangcheng, Huangbaishan, 200 m, 14.VII.1999, Xiaocheng Shen &

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 27 Yingdang Ren (CAU). Distribution. China (Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Jiangxi, Sichuan, Zhejiang). Remarks. Protohermes gutianensis appears to be closely related to P. acutatus by the absence of the apical round markings of both wings, but it can be easily separated from P. acutatus by having the head with a pair of slender blackish vittae and the male ninth sternum with deep suboblong apical incision, both of which are unique in the P. c o st al is group.

FIGURES 75–81. Protohermes gutianensis Yang & Yang. 75. Male head and pronotum, dorsal view. 76. Male genitalia, dorsal view. 77. Male genitalia, ventral view. 78. Male ninth gonostylus, caudal view. 79. Male tenth sternum, ventral view. 80. Female genitalia, lateral view. 81. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes hunanensis Yang & Yang (Figs. 14, 82–89)

Protohermes hunanensis Yang & Yang, 1992. Iconography of forest insects in Hunan, China, p. 641. Type locality: Hunan (Yongshun).

Diagnosis. Head entirely black near lateral margins; antenna and mandibles entirely black; pronotum with one pair of blackish vittae on lateral margins; wings dark with distinct whitish markings. Body length 33–35 mm (male) and 35–38 mm (female); forewing length 35–37 mm (male) and 37–39 mm (female), hindwing length 32–33 mm (male) and 33–36 mm (female). Head (Fig. 82) yellow without post-ocular spine, dorsally entirely black near lateral margins. Compound

28 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. eyes brown; ocelli yellow, darkly margined medially. Antennae black. Mouthparts mostly yellow with black mandible. Occiput yellow with one pair of blackish markings on lateral margins. Prothorax (Fig. 82) yellow; pronotum with one pair of blackish vittae on lateral margins. Meso- and metathorax brown, each dorsally with one pair of blackish spots on lateral sides. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs black with dense, pale yellow, short setae, tarsal claws reddish brown. Forewings (Fig. 14) dark, with 1 large and 1 small whitish markings at base, 2 large and 4 small whitish markings at middle, and 1 distinctly enlarged round whitish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings dark except basal half hyaline, respectively with 1 large round whitish marking at middle and apical 1/3. Veins dark brown, except pale yellow veins in whitish markings and base of hindwings. Rs 8 to 9-branched, last branch mostly bifurcate; 10–13 crossveins between R1 and Rs; M1+2 5 to 7- branched, M3+4 3-branched; 1A 3-branched.

FIGURES 82–89. Protohermes hunanensis Yang & Yang. 82. Male head and pronotum, dorsal view. 83. Male genitalia, dorsal view. 84. Male ninth sternum, ventral view. 85. Male ninth gonostylus, caudal view. 86. Male tenth tergum, ventral view. 87. Male tenth sternum, ventral view. 88. Female genitalia, lateral view. 89. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen gray with short, yellow setae. Male ninth tergum (Fig. 83) subtrapezoidal, with parabolic anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 84) broad; posterior margin with extremely shallow trapezoidal incision, forming two broad roundly produced processes; central portion inflated. Ninth gonostylus (Fig. 85) unguiform and strongly curved dorsomedially. Tenth tergum (Fig. 86) short and subcylindrical, ventrally with one feebly developed tufted tubercle anteromedially; posterolateral

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 29 corner distinctly and roundly produced; tip slightly incised and densely setose. Tenth sternum (Fig. 87) arched, anterior margin slightly elevated, and posterior incision indistinct; lateral lobes slender digitiform and slightly incurved. Female eighth sternum (Figs. 88–89) strongly sclerotized and subtrapezoidal in lateral view, with truncate posterior margin; ninth segment (Fig. 88) laterally inflated in one pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one triangular upper and one subquadrate lower lobe. Material examined. Holotype %, CHINA: Hunan, Yongshun, 20.VI.1981 (CAU). CHINA : 2 &, Guangxi, Luoxiang, 200 m, 15.V.1999, Xuezhong Zhang (IZCAS); &, Guangxi, Luoxiang, 400 m, 15.V.1999, Wenzhu Li (IZCAS); %, Guangxi, Napo, Defu, 1350 m, 9.VI.2000, Wenzhu Li (IZCAS); &, Guangdong, Dinghushan, 28.IV.1965, Youwei Zhang (IZCAS). Distribution. China (Guangdong, Guangxi, Hunan). Remarks. Protohermes hunanensis is the only species in P. co s ta li s group with dark body coloration and is similar to species of Neurhermes in this respect. The genitalia of P. hunanensis is similar to that of P. fujianensis in having the more trapezoidal male tenth tergum in dorsal view.

Protohermes lii sp. nov. (Figs. 15, 90–94)

Diagnosis. Head without dark markings; pronotum with two small blackish bivittate on lateral sides; male ninth sternum with wide, shallow posterior incision; male tenth tergum with feebly developed ventral tufted tubercle. Body length 23–25 mm (male); forewing length 37–39 mm (male), hindwing length 34–35 mm (male). Head (Fig. 90) yellow, without post-ocular spine; vertex without markings. Compound eyes pale brown; ocelli yellow, darkly margined medially. Antennae dark, scapes and pedicel yellow. Mouthparts yellow; mandible distally dark. Prothorax yellow; pronotum (Fig. 90) with two pairs of widely separated small blackish markings on each side. Meso- and metathorax pale yellow. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs yellow with dense yellowish short setae; wide apices of fore and mid tibiae and 2nd–5th tarsi dark; tarsal claws reddish brown. Forewings (Fig. 15) pale smoky brown, with 1 large yellowish marking at base, 3 mostly connected large and 5–7 small yellowish markings on crossveins between R1 and Rs at middle, and 1 round yellowish marking on apical 1/3; costal cellules with distinct brown stripes. Hindwings much paler than forewings, respectively with 1 round yellowish marking at middle and apical 1/3. Veins brown except veins in yellowish markings and basal half of hindwings pale yellow. Rs 9 to 10-branched, last branch bifurcate; 10–13 crossveins between R1 and Rs; M1+2 6 to 7-branched, M3+4 3-branched; 1A 3-branched. Abdomen yellow with short, yellow setae. Male ninth tergum (Fig. 91) subtrapezoidal, with arched anterior margin and sightly incised posterior margin. Ninth sternum (Fig. 92) broad with lateral margin slightly curved near anterior 1/4, posterior margin with broad, shallow incision, forming two acute-angled posterior processes; central portion inflated. Ninth gonostylus (Fig. 93) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 91–92) short, subcylindrical, ventrally with one feebly developed tufted tubercle near posteromedial corner; posterolateral corner slightly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 94) slightly arched, anteriorly slightly elevated at middle, posterior margin with extremely small V-shaped median incision and a pair of small denticles; lateral lobes digitiform, with apex strongly narrowed and incurved. Female unknown. Type material. Holotype %, CHINA: Guangxi, Fangcheng, Dongzhong, 360–700 m, 9.IV.2002, Aimin Shi (HBU). Paratypes %, CHINA: same data as holotype (HBU); %, CHINA: Guangxi, Shangsi, Pinglongshan,

30 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. 350–500 m, 6.IV.2002, Aimin Shi (HBU); %, CHINA: Guangxi, Shiwandashan, VIII.2002, Ding Yang (CAU). Etymology. The specific epithet ‘lii’ refers to Prof. Fasheng Li, who is a famous entomologist on Psocop- tera and Hemiptera at the China Agricultural University. We thank him for his kind help with our studies to collect numerous megalopteran specimens during the past thirty years. Distributions. China (Guangxi). Remarks. The new species somewhat resembles P. basiflavus, but can be easily separated by the smaller body size, the widely separated, small prothoracic black markings, the feebly developed ventral tufted turbercle of the male tenth tergum, and the male tenth sternum with the posterior margin bearing a pair of small denticles. In P. basiflavus, the body size and the prothoracic markings are much larger, the ventral tufted turbercle of the male tenth tergum is well developed, and the male tenth sternum posteriorly bears a pair of distinctly produced processes.

FIGURES 90–94. Protohermes lii, sp. nov. 90. Male head and pronotum, dorsal view. 91. Male genitalia, dorsal view. 92. Male genitalia, ventral view. 93. Male ninth gonostylus, caudal view. 94. Male tenth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes niger Yang & Yang (Figs. 16, 95–100)

Protohermes niger Yang & Yang, 1988. Zool. Res. 9 (1): 51. Type locality: Yunnan (Fengqing, Lincang).

Diagnosis. Head with one pair of blackish markings on lateral margins; pronotum dark, with several indistinct yellowish brown markings; wings grayish brown with several whitish markings. Body length 27–28 mm; forewing length 35–36 mm, hindwing length 32–33 mm. Head (Fig. 95) yellow or yellowish brown with small post-ocular spine, vertex with one pair of blackish markings on lateral margins. Compound eyes brown; ocelli yellow, darkly margined medially. Median ocellus slightly transverse; posterior pair widely apart, but distance between them slightly shorter than that between

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 31 antennal bases. Antennae black with scapes and pedicel yellowish brown. Mouthparts yellow; mandible distally black. Prothorax (Fig. 95) dark; pronotum with yellowish brown anterior margin, and with one slender yellow brown spot near each anterolateral corner and one long hook-shaped yellow brown marking following the former spot; central marking cuneiform, yellowish brown, extending to middle. Meso- and metathorax brown, each dorsally with one pair of brownish spots on lateral sides. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs pale brown with dense pale yellowish short setae, tarsal claws reddish brown. Forew- ings (Fig. 16) grayish brown, with 1 large and 2 small whitish marking at base, 6–7 whitish markings at middle and 1 round whitish marking on apical 1/3; costal area brown. Hindwings grayish brown, with 1 indistinct round whitish marking at base, 1 indistinct round whitish marking at middle and 1 distinct round whitish marking on apical 1/3. Veins dark, except pale yellow on whitish markings. Rs 7 to 8-branched, last branch bifurcate; 6–9 crossveins between R1 and Rs; M1+2 4-branched, M3+4 2-branched; 1A 3-branched.

FIGURES 95–100. Protohermes niger Yang & Yang. 95. Male head and pronotum, dorsal view. 96. Male genitalia, dorsal view. 97. Male ninth sternum, ventral view. 98. Male ninth gonostylus, caudal view. 99. Male tenth tergum, ventral view. 100. Male tenth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen brown with yellowish short setae. Male ninth tergum (Fig. 96) subquadrate, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 97) broad, posterior margin with shallow trapezoidal incision, forming two broad acute-angled triangular processes; central portion feebly inflated. Ninth gonostylus (Fig. 98) unguiform and strongly curved dorsomedially. Tenth tergum (Fig. 99) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner not produced; tip slightly incised and densely setose. Tenth sternum (Fig. 100) arched with ante-

32 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. rior margin feebly elevated; posterior incision triangular and extremely small; lateral lobes thick digitiform and slightly incurved. Female unknown. Material examined. Holotype %, CHINA: Yunnan, Fengqing, Sanchahe, 1200 m, 24.VII.1980 (CAU). Paratypes: %, CHINA: same data as holotype (CAU); %, CHINA: Yunnan, Lincang, 1750 m, 4.VII.1980, Qiaosheng Li (CAU); %, CHINA: Yunnan, Fengqing, Sanchahe, 1400 m, 20.VII.1980 (CAU). %, CHINA: Yunnan, Tengchong, Houqiao, 1700 m, 2.VI.2005, Hao Huang (CAU); %, CHINA: Yunnan, Tengchong, Jietou, 1500 m, 17.V.2006, Ping Zhao (CAU); %, CHINA: Yunnan, Nanjiangonglang, 9.VII.1981, Dazhi Dong (KIZCAS); 2 %, CHINA: Yunnan, Baoshan, Wayao, 7.VI.1983, Dazhi Dong (KIZCAS); %, CHINA: Yunnan, Lincang, 20.VI.1979, Yunzhen Wang (KIZCAS); %, CHINA: Yunnan, Lincang, 4.VII.1981, Dazhi Dong (KIZCAS); %, CHINA: Yunnan, Lincang, 5.VII.1981, Pingzhi Yang (KIZCAS); 2 %, CHINA: Lincang, Yongde, 10–23.VII.1981, Wenguo Yang (KIZCAS). Distribution. China (Yunnan). Remarks. This species is distinguished from the other species of the P. c os ta l is group by the distinct blackish pronotum.

Protohermes orientalis sp. nov. (Figs. 17, 101–107)

Diagnosis: Head with three blackish markings on each side of vetex; antenna entirely black; pronotum with two blackish markings on each side; legs almost dark; male ninth sternum with extremely shallow trapezoidal posterior incision. Body length 25–32 mm (male) and 32–35 mm (female); forewing length 31–39 mm (male) and 41–45 mm (female), hindwing length 30–31 mm (male) and 36–38 mm (female). Head (Fig. 101) yellowish brown, without post-ocular spine; vertex with three blackish markings on each side, anterior marking large and subquadrate, posterolateral marking cuneiform, posteromedial marking punctuate. Compound eyes brown; ocelli yellowish brown, darkly margined medially. Antennae black. Mouthparts yellowish brown; maxillary palpus and labial palpus dark brown, mandible with distal half black. Prothorax (Fig. 101) yellowish brown; pronotum with two blackish markings on each side. Meso- and metathorax dark, dorsally with each center pale yellow. Thoracic pilosity yellow, much longer on meso- and meta thorax. Legs dark with dense yellowish short setae; fore and mid legs with trochanters and dorsal surface of femora black; tarsal claws reddish brown. Forewings (Fig. 17) pale smoky brown, with 1 yellowish marking at base, 3–4 yellowish markings at middle and 1 round yellowish markings on apical 1/3; costal cellules with brownish stripes; veins dark except veins in yellowish markings pale yellow. Hindwings much paler than forewings, respectively with 1 yellowish marking at middle and apical 1/3; veins dark, except veins in yellowish markings and at base pale yellow. Rs 9 to 10-branched, last branch bifurcate; 9–11 crossveins between R1 and Rs; M1+2 6 to 7-branched, M3+4 2 to 3-branched; 1A 3-branched. Abdomen dark with yellowish short setae. Male ninth tergum (Fig. 102) subquadrate, with arched anterior margin and truncate posterior margin. Ninth sternum (Fig. 103) subquadrate, posterior margin with extremely shallow incision, leaving a pair of small widely separated subtriangular posterior processes; central portion inflated. Ninth gonostylus (Fig. 104) unguiform and slightly curved dorsomedially. Tenth tergum (Fig. 102– 103) short and subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 105) arched, anteriorly slightly elevated at middle; posterior incision extremely small and V-shaped; lateral lobes elongated and digitiform, distally incurved. Female eighth sternum (Figs. 106–107) strongly sclerotized, subtriangular in lateral view, and subtrapezoidal in ventral view, posterior margin slightly incised,

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 33 forming pair of obtuse processes; ninth segment (Fig. 106) laterally inflated in one pair of small sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one somewhat long digitiform upper and one short lower lobe. Type materials. Holotype %, CHINA: Fujian, Jiangle, Longqishan, 11.V.1999, Jian Yao (IZCAS). Paratypes %, CHINA: same data as holotype (IZCAS); %, CHINA: Fujian, Jiangle, Longqishan, 28.IV.1999, Wanyu Zhang (IZCAS); &, CHINA: Fujian, Jiangle, Longqishan, 11.V.1999, Chunmei Huang (IZCAS); &, CHINA: Fujian, Jiangle, Longqishan, 11.V.1999, Runzhi Zhang (IZCAS); %, CHINA: Fujian, Jiangle, Long- qishan, 15.V.1999, Jian Yao (IZCAS); &, CHINA: Fujian, Jiangle, Longqishan, 16.V.1999, Jian Yao (IZCAS); &, CHINA: Guangxi, Jinxiu, Luoxiang, 200 m, 14.V.1999, Xuezhong Zhang (IZCAS); % &, CHINA: Guangxi, Maoershan, Tongren, 350 m, 2.V.2004, Ding Yang (CAU); 6 % 10 &, CHINA: Jiangxi, Jiulianshan, 400 m, VII.1986, Zuyao Liu (SEMCAS). Etymology. The specific epithet ‘orientalis’ refers to the oriental distribution of this species. Distribution. China (Fujian, Guangxi, Jiangxi). Remarks. The new species appears to be closely related to P. acutatus in having a similar appearance, but can be easily separated by the male ninth sternum which has extremely shallow posterior incision. In P. acutatus, the male ninth sternum is deeply incised posteriorly.

FIGURES 101–107. Protohermes orientalis, sp. nov. 101. Male head and pronotum, dorsal view. 102. Male genitalia, dorsal view. 103. Male genitalia, ventral view. 104. Male ninth gonostylus, caudal view. 105. Male tenth sternum, ventral view. 106. Female genitalia, lateral view. 107. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes similis Yang & Yang, 1988. Zool. Res. 9 (1): 47. Type locality: Sichuan (Ya-an, Emeishan), Yunnan (Yan- jin). Protohermes sichuanensis Yang & Yang, 1997. J. China Agric. Univ. 2 (5): 31. Type locality: Sichuan (Emeishan). syn. nov.

Diagnosis. Head with one large blackish marking on each side; pronotum with two pairs of blackish markings on each side; forewings with distinct brownish stripes on costal cellules; male ninth sternum with trapezoidal posterior incision; male tenth sternum with extremely small V-shaped posterior incision and short digitiform lateral lobes. Body length 30–32 mm (male) and 40–45 mm (female); forewing length 42–44 mm (male) and 50–52 mm (female), hindwing length 39–41 mm (male) and 45–46 mm (female). Head (Fig. 108) pale yellowish brown to reddish brown without post-ocular spine; vertex with one large blackish marking on each side, occasionally separated into three markings. Compound eyes brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel brown. Mouthparts yellowish brown; mandible with distal half black. Occiput yellowish brown with one pair of blackish markings on lateral sides. Prothorax (Fig. 108) yellowish brown or entirely brown; pronotum with two blackish markings on each side. Meso- and metathorax yellowish brown, each dorsally with one pair of brownish markings on sides. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs yellow to brown with dense yellowish short setae; tibiae and tarsi black; tarsal claws reddish brown. Forewings (Fig. 18) pale smoky brown, with 1– 3 yellowish markings at base, 4–8 yellowish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings much paler than forewings, respectively with 1 round yellowish marking at middle and apical 1/3. Veins pale brown, except costal crossveins pale yellow and veins in yellowish markings yellow. Rs 9 to 10-branched, last branch trifurcate; 10–13 crossveins between R1 and Rs; M1+2 5 to 7-branched, M3+4 4-branched; 1A 3-branched. Abdomen brown with short, yellow setae. Male ninth tergum (Fig. 109) subtrapezoidal, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 110) broad with arched lateral margins; posterior margin with trapezoidal incision, forming two broad acute-angled triangular processes; central portion inflated. Ninth gonostylus (Fig. 111) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 108–109) short and subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; posterolateral corner slightly produced; tip slightly incised and densely setose. Tenth sternum (Fig. 112) arched, anteriorly slightly elevated at middle; posterior incision V-shaped; lateral lobes short digitiform and slightly incurved. Female eighth sternum (Fig. 113–114) strongly sclerotized and subtrapezoidal in lateral view, posterior margin apparently elevated with shallow median incision, forming pair of obtuse processes; ninth segment (Fig. 113) laterally inflated one pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one thick digitiform upper and one round lower lobe. Material examined. Holotype %, CHINA: Sichuan, Ya-an, VIII.1984 (CAU). Allotype &, CHINA: same data as holotype (CAU). Paratypes, %, CHINA: same data as holotype (CAU); &, CHINA: Sichuan, Emeis- han, Qingyinge, 22.VII.1962, Zhenghong Yu (CAU); 2 &, CHINA: Yunnan, Yanjin, 800 m, 12.VI.1980, Zongli Hu (CAU). %, CHINA: Sichuan, Emeishan, Wanniansi, 23.V.1962, Zhaorong Chen & Yizheng Wang (CAU); &, CHINA: Sichuan, Emeishan, Wanniansi, 28.VII.1962, Zhaorong Chen (CAU); &, CHINA: Sichuan, Qingchengshan, Tianshidong, 1000 m, 4.VI.1979, Ping Gao (IZCAS); &, CHINA: Sichuan, Qingchengshan, 1000 m, 5.VI.1979, Jinwen Gao (IZCAS); &, CHINA: Sichuan, Emeishan, 710 m, 20.VI.1979, Ping Gao (IZCAS); %, CHINA: Sichuan, Emeishan, Qingyinge, 800–1000 m, 30.VI.1957 (IZCAS); &, CHINA: Sichuan, Emeishan, 710 m, 20.VI.1979, Ping Gao (IZCAS); &, CHINA: Sichuan, Ya-

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 35 an, 1973 (CAU); &, CHINA: Sichuan, Ya-an, 1974 (CAU). Distribution. China (Guizhou, Sichuan, Yunnan). Remarks. Protohermes similis appears to have a close affinity with P. acutatus in having similar general structure of the male genitalia and similar pattern of markings on head, prothorax and wings. But this species is distinguished from P. acutatus by having the forewings with distinct brown stripes on the costal cellules and round yellowish markings at the apical 1/3, while in P. acutatus the forewings are much paler, with indistinct stripes on the costal cellules and the apical portions of the forewings. Protohermes sichuanensis was established by Yang and Yang (1997) according to one male and one female specimen. After examination to the type series of P. sichuanensis, no distinct differences from P. similis were found. Contrary to what was stated in the original description, P. sichuanensis has the trapezoidal apical incision on the male ninth sternum, which is the same as P. si m il is . Therefore, P. sichuanensis is undoubtedly a new synonym of P. similis.

FIGURES 108–114. Protohermes similis Yang & Yang. 108. Male head and pronotum, dorsal view. 109. Male genitalia, dorsal view. 110. Male genitalia, ventral view. 111. Male ninth gonostylus, caudal view. 112. Male tenth sternum, ventral view. 113. Female genitalia, lateral view. 114. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes sinensis Yang & Yang (Figs. 19, 115–122)

Protohermes sinensis Yang & Yang, 1992. Iconogrphy of forest insects in Hunan China, p. 640. Type locality: Hunan (Cili), Zhejiang (Lishui).

36 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. Diagnosis. Head with one transverse dark marking anterior to ocellar triangle and three blackish markings on each lateral side of vertex; pronotum with one dark vitta on each lateral side; forewing costal region entirely hyaline; male ninth sternum with wide shallow V-shaped posterior incision. Body length 20–25mm (male) and 29–40 mm (female); forewing length 35–37 mm (male) and 45–47 mm (female), hindwing length 31–33 mm (male) and 39–43 mm (female). Head (Fig. 115) pale yellow to brown without post-ocular spine; a transverse dark marking occurs anterior to ocellar triangle; vertex with three blackish markings on each side, anterior marking large and subquadrate, posterolateral marking cuneiform, posteromedial marking punctuate. Compound eyes brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel pale yellow or brown. Mouthparts yellowish brown; mandible with distal half black.

FIGURES 115–122. Protohermes sinensis Yang & Yang. 115. Male head and pronotum, dorsal view. 116. Male genitalia, dorsal view. 117. Male ninth sternum, ventral view. 118. Male ninth gonostylus, caudal view. 119. Male tenth tergum, ventral view. 120. Male tenth sternum, ventral view. 121. Female genitalia, lateral view. 122. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Thorax pale yellow to brown. Pronotum (Fig. 115) with one dark vitta on each side. Meso- and metanota each with one pair of blackish markings laterally. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs dark with dense yellowish short setae; coxae, trochanters and basal tibiae yellow, sometimes tibiae entirely brown; tarsal claws reddish brown. Forewings (Fig. 19) pale smoky brown, with 1 yellowish marking at base, 3–4 suboval yellowish markings at middle and 1 round yellowish marking on apical 1/3; veins brown, except veins in yellowish markings yellow. Hindwings much paler than forewings, with 1 round yellowish marking at middle and apical 1/3; veins brown, except veins in yellowish markings and basal longi-

Rs; M1+2 5 to 7-branched, M3+4 2 to 3-branched; 1A 3-branched. Abdomen brown with yellowish short setae. Male ninth tergum (Fig. 116) subquadrate, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 117) subquadrate with straight lateral margin, apical margin with shallow V-shaped incision, forming two small acutely angled apical processes; central portion inflated. Ninth gonostylus (Fig. 118) unguiform and slightly curved dorsomedially. Tenth tergum (Fig. 119) short and subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; posterolateral corner not produced; tip slightly incised and densely setose. Tenth sternum (Fig. 120) arched, anterior slightly elevated at middle; posterior incision extremely small and V-shaped; lateral lobes digitiform and slightly incurved. Female eighth sternum (Figs. 121–122) strongly sclerotized and subtrapezoidal in lateral view, posterior margin elevated with shallow median incision, forming pair of obtuse processes; ninth segment (Fig. 121) laterally inflated as a pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with a digitiform process on tip; tenth tergum short, with posterior margin medially incised, leaving one somewhat long digitiform upper and one short round lower lobe. Material examined. Holotype %, CHINA: Zhejiang, Lishui, 2.V.1978 (CAU). Allotype &, CHINA: same data as holotype (CAU). Paratypes 7 % 14 &, CHINA: same data as holotype (CAU); &, CHINA: Hunan, Cili, 20.V.1981(CAU). 2 &, CHINA: Zhejiang, Moganshan, 22–23.V.1936 (IZCAS); %, CHINA: Henan, Luoshan, Lingshan, 300 m, 21–25.V.2000, Xiaocheng Shen & Yingdang Ren (CAU); &, CHINA: Henan, Huanglian- shu, 600 m, 5.VI.2000(CAU). Distribution. China (Henan, Hunan, Zhejiang). Remarks. This species resembles P. xanthodes in having similar wing coloration, but it can be easily separated from P. xanthodes by the subcylindrical male tenth tergum and the presence of the sac-like lobes in the female genitalia. The male ninth sternum of P. si ne n si s is distinguished from the other species of the P. c o st a - lis group in having extremely shallow and wide V-shaped posterior incision.

Protohermes stigmosus sp. nov. (Figs. 20, 123–127)

Diagnosis. Head without dark markings; pronotum with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings at posterolateral corner; male ninth sternum has a wide U-shaped posterior incision, leaving two short acutely angled digitiform processes; tenth sternum with posterior incision strongly developed, leaving two large papillate processes with slender digitiformed tip, lateral lobes short and digitiform, proximally strongly inflated and narrowing toward tip. Body length 25 mm; forewing length 38 mm, hindwing length 35 mm. Head (Fig. 123) yellow with short and obtuse post-ocular spine; vertex without markings. Compound eyes brown; ocelli yellowish brown, darkly margined medially. Antennae black, with scapes and pedicel yellow. Mouthparts yellow; mandible with distal half black. Prothorax (Fig. 123) yellow; pronotum with two pairs of blackish markings at anterolateral corner and three pairs of blackish markings at posterolateral corner. Meso- and metathorax pale yellowish brown. Tho- racic pilosity yellow, much longer on meso- and metathorax. Legs yellow with dense yellowish short setae; distal half of fore tibiae and all fore tarsi black, middle and hind tarsi with 2nd–5th tarsomeres black; tarsal claws pale reddish brown. Forewings (Fig. 20) pale grayish brown except anal region entirely hyaline, with 1 subtriangular yellowish marking near base,1 broad and 1 narrow yellowish markings at middle, 1 round and 1 narrow yellowish markings on apical 1/3 and several extremely small yellowish markings on crossveins at apical portion; costal cellules with distinct grayish brown stripes. Hindwings entirely hyaline. Veins pale yellow except veins in dark regions of forewings and apical portion of hindwings pale brown. Rs 8-branched, last

38 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. branch bifurcate; 9–12 crossveins between R1 and Rs; M1+2 4-branched, M3+4 2-branched; 1A 3-branched. Abdomen pale yellowish brown with yellowish short setae. Male ninth tergum (Fig. 124) much wider posteriorly, with deep U-shaped anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 125) suboblong with central portion apparently inflated; posterior margin with wide U-shaped incision, forming two short, acutely angled digitiform processes. Ninth gonostylus (Fig. 126) unguiform and elongated, slightly curved dorsomedially. Tenth tergum (Figs. 124–125) short and subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; tip slightly incised and densely setose. Tenth sternum (Fig. 127) arched, produced anteromedially; posterior incision strongly developed, forming two large median papillate processes, lateral lobes slender digitiform; lateral arms short and digitiform, proximally inflated and strongly narrowing toward tip. Female unknown. Type material. Holotype %, CHINA: Yunnan, Xishuangbanna, 31.iii.2004, reared from larva, Liang Tang (SNU). Etymology. The specific epithet ‘stigmosus’ refers to the complex pattern of the markings on the pronotum and the forewings of this species. Distribution. China (Yunnan). Remarks. This new species is distinguished from all the other members in the P. c o st al is group based on the ten small dark, lateral spots on the pronotum, by the male ninth sternum with a wide, U-shaped posterior incision, and by the male tenth sternum with the abnormal posterior lobes. However, the more complicated wing patterns somewhat show the close affinity between P. st ig m o s us and P. triangulatus.

FIGURES 123–127. Protohermes stigmosus, sp. nov. 123. Male head and pronotum, dorsal view. 124. Male genitalia, dorsal view. 125. Male genitalia, ventral view. 126. Male ninth gonostylus, caudal view. 127. Male tenth sternum, ventral view. Scale lines = 1.0 mm.

Protohermes triangulatus sp. nov. (Figs. 21, 128–134)

Diagnosis. Head with two small blackish markings on each side of vertex; pronotum with two large blackish markings on each side; male ninth sternum with deep V-shaped posterior incision; tenth tergum with an

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 39 exceptionally well developed tufted tubercle at posteromedial corner. Body length 27–34 mm (male) and 35–41 mm (female); forewing length 37–41 mm (male) and 46–50 mm (female), hindwing length 32–36 mm (male) and 39–44 mm (female). Head (Fig. 128) pale yellow without post-ocular spine; vertex with two small blackish markings on each side, lateral marking cuneiform, medial marking punctuate, and sometimes markings almost absent. Com- pound eyes brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel yellow. Mouthparts yellow; mandible with distal 1/3 black. Occiput with one pair of blackish markings on lateral sides. Thorax yellow. Pronotum (Fig. 128) with two small blackish markings on each side. Meso- and metanota each with one pair of brownish markings on lateral sides. Thoracic pilosity yellow, much longer on meso- and meta thorax. Legs yellow with dense yellowish short setae; tibiae and tarsi mostly dark, and sometimes only 2nd–5th mid and hind tarsomeres dark; tarsal claws reddish brown. Forewings (Fig. 21) light smoky brown, with 1 whitish marking at base, 4–5 mostly fused whitish markings at middle and 1 round whitish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings mostly hyaline, respectively with 1 indistinct round whitish marking on apical 1/3. Veins yellowish brown, with costal crossveins much paler. Rs 9- branched, last branch bifurcate or trifurcate; 9–10 crossveins between R1 and Rs; M1+2 4 to 6-branched, M3+4 2- branched; 1A 3-branched.

FIGURES 128–134. Protohermes triangulatus, sp. nov. 128. Male head and pronotum, dorsal view. 129. Male genitalia, dorsal view. 130. Male genitalia, ventral view. 131. Male ninth gonostylus, caudal view. 132. Male tenth sternum, ventral view. 133. Female genitalia, lateral view. 134. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen pale brown with yellowish short setae. Male ninth tergum (Fig. 129) subquadrate, with arched anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 130) broad with nearly straight lateral margin; posterior margin with deep V-shaped incision; central portion inflated. Ninth gonostylus (Fig.

40 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. 131) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 129–130) short and subcylindrical, ventrally with one well developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 132) arched, anteriorly slightly elevated at middle; posterior incision V-shaped and moderately developed; lateral lobes digitiform and slightly incurved. Female eighth sternum (Figs. 133–134) strongly sclerotized and subtrapezoidal in lateral view, with truncate posterior margin; ninth segment (Fig. 133) laterally inflated in one pair of small suboval lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one round lower lobe. Material examined. Holotype %, CHINA: Yunnan, Xinping, 19.VII.1980 (CAU). Paratypes &, CHINA: Yunnan, Shuangjiang, 858 m, 5.VI.1980, Wang Zhang (CAU); %, CHINA: Yunnan, Yongping, 1700 m, 26.V.1980 (CAU); %, CHINA: Yunnan, Anning, 2000 m, 17.VII.1980 (CAU); %, CHINA: Yunnan, Fengqing, Songxiangchang, 1500 m, 3.VII.1980, Fu Zhang (CAU); % &, CHINA: Yunnan, Gejiu, 11.VIII.1980, Guoxiu Li (CAU); 1 without genitalia, CHINA: Yunnan, Yimen (CAU); %, CHINA: Yunnan, Jingdong, 1170 m, 30.V.1956 (IZCAS); &, CHINA: Yunnan, Jingdong, 1170–1300 m, 30.V.1956 (IZCAS); %, CHINA: Yunnan, Jingdong, 1170 m, 2.VI.1956 (IZCAS); %, CHINA: Yunnan, Lancang, 1000 m, 3.III.1957, Shuyong Wang (IZCAS); &, CHINA: Yunnan, Xishuangbanna, Yunjinghong, 2.VII.1958, Chuanlong Li (IZCAS). Etymology. The specific epithet ‘triangulatus’ refers to the triangular or V-shaped posterior incision of the male ninth sternum in the new species. Distribution. China (Yunnan). Remarks. The new species appears to be closely related to P. arunachalensis, but can be easily separated by the V-shaped posterior incision of the male ninth sternum. In P. arunachalensis, the male ninth sternum has a U-shaped posterior incision.

Protohermes yangi sp. nov. (Figs. 22, 135–141)

Diagnosis. Head with two blackish markings on each side of vertex; pronotum bivittate black; wings grayish brown with distinct whitish markings; male ninth sternum with deep V-shaped posterior incision; male tenth tergum strongly and roundly produced distolaterally. Body length 31–34 mm (male) and 35–40 mm (female); forewing length 37–40 mm (male) and 38–45 mm (female), hindwing length 33–35 mm (male) and 35–41 mm (female). Head (Fig. 135) yellowish brown without post-ocular spine; vertex with two blackish markings on each side, lateral marking band-like, medial marking punctuate. Compound eyes grayish brown; ocelli yellow, darkly margined medially. Antennae black, scapes and pedicel brown with tip and basal half pale yellow. Mouthparts yellowish brown; distal three joints of maxillary palpus and distal three joints of labial palpus brown, mandible with distal half dark. Occiput yellowish brown with one pair of blackish markings on lateral sides. Prothorax (Fig. 135) yellowish brown; pronotum with one blackish vitta on each side, vittae sometimes narrowed medially. Meso- and metathorax pale brown, and mesonotum dorsally with pale yellow center. Tho- racic pilosity yellow, much longer on meso- and metathorax. Legs pale yellow to brown with dense yellowish short setae; wide apical portion of tibiae and tarsi black; tarsal claws reddish brown. Forewings (Fig. 22) grayish brown, with 1 subtriangular whitish marking at base, 2 large and 5–6 small whitish markings at middle and 1 extremely enlarged round whitish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings pale brown with basal half mostly hyaline, with 1 round whitish marking on apical 1/3. Veins yellowish brown, except veins on apical half pale brown. Rs 8 to 10-branched, last branch bifurcate or trifurcate;

9–11 crossveins between R1 and Rs; M1+2 6 to 7-branched, M3+4 2 to 3-branched; 1A 3-branched.

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 41 FIGURES 135–141. Protohermes yangi, sp. nov. 135. Male head and pronotum, dorsal view. 136. Male genitalia, dorsal view. 137. Male genitalia, ventral view. 138. Male ninth gonostylus, caudal view. 139. Male tenth sternum, ventral view. 140. Female genitalia, lateral view. 141. Female eighth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen dark with yellowish short setae. Male ninth tergum (Fig. 136) subquadrate, with trapezoidal anterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 137) broad with straight lateral margin; posterior margin with deep V-shaped incision, forming two large triangular processes; central portion inflated. Ninth gonostylus (Fig. 138) unguiform and strongly curved dorsomedially. Tenth tergum (Figs. 136– 137) short and subcylindrical, ventrally with one feebly developed tufted tubercle at posteromedial corner; posterolateral corner distinctly and roundly produced; tip densely setose and sometimes slightly incised. Tenth sternum (Fig. 139) arched, anteriorly with one subtriangular median process; posterior incision small and V- shaped; lateral lobes digitiform and slightly incurved. Female eighth sternum (Figs. 140–141) strongly sclerotized and subtriangular in lateral view, posterior margin elevated without incision, forming a pair of triangular processes; ninth segment (Fig. 140) laterally inflated forming a pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one round lower lobe. Type materials. Holotype %, VIETNAM: Mt. Pia Oac, 14.V.1998, F. Hayashi (CAU). Paratypes: 3 %, CHINA: Guangxi, Tianlin, Langping, 1300 m, 2.VI.2002, Jianwen Liu (CAU); 2 %, CHINA: Guangxi, Tian- lin, Langping, 1300 m, 1.VI.2002, Jianwen Liu (CAU); %, CHINA: Guangxi, Linaoshan, 1400 m, 28.V.2002, Jianwen Liu (CAU); %, CHINA: Guangxi, Linaoshan, 1400 m, 28.V.2002, Guofang Jiang (CAU); %, CHINA: Guangxi, Tianlin, Laoshan Forestry Center, 1400 m, 5.VI.2002, Guofang Jiang (CAU); % &, CHINA: Gaungxi, Tianlin, Yaojiawan, 1200–1400 m, 1.VI.2002, Xiujuan Yang (HBU); &, CHINA, Guangxi, Tian-e, Buliuhe, 6.VIII.2002, Ding Yang (CAU); &, CHINA: Guizhou, Pingtang, Yenanzhou (CAU). Etymology. The specific epithet ‘yangi’ refers to the famous entomologist Prof. Chikun Yang, who passed away in 2006, for his great contributions to the systematics of the Chinese Megaloptera.

42 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. Distribution. China (Guangxi, Guizhou); Vietnam (Mt. Pia Oac). Remarks. This new species might have a close affinity with P. hunanensis and P. fujianensis due to the similar wing pattern with extremely enlarged round whitish marking on the apical 1/3 of both wings. How- ever, the male ninth sternum of the new species bears a deep V-shaped posterior incision. Alternatively, the posterior incision of the male ninth sternum is trapezoidal in both P. hunanensis and P. fujianensis. The new species also somewhat resembles P. triangulatus in having the similar V-shaped posterior incision of the male ninth sternum, but it can be distinguished by the male tenth tergum ventrally with feebly developed tubercle. In P. triangulatus, the ventral tubercle is well developed.

Protohermes yunnanensis Yang & Yang (Figs. 23, 142–148)

Protohermes yunnanensis Yang & Yang, 1988. Zool. Res. 9 (1): 48. Type locality: Yunnan (Yunxian, Shuangjiang, Jing- hong, Yongping, Anning, Xinping, Fengqing, Yimen, Tengchong, Gejiu).

Diagnosis. Head immaculate; pronotum with two narrow, dark markings on each side; male ninth sternum with truncate posterior margin. Body length 30–31 mm (male) and 38–43 mm (female); forewing length 31–39 mm (male) and 48–50 mm (female), hindwing length 35–36 mm (male) and 40–43 mm (female). Head (Fig. 142) yellow, without post-ocular spine; vertex without markings. Compound eyes brown; ocelli yellow, darkly margined medially. Anttenae black with scapes and pedicel yellow. Mouthparts yellow or yellowish brown; mandible with distal half black. Thorax yellow. Pronotum (Fig. 142) with two narrow blackish markings on each side. Meso- and metanota each with posterolateral corners pale brown. Thoracic pilosity yellow, much longer on meso- and metathorax. Legs pale yellow with dense yellowish short setae; apical 1/3 of fore tibiae, fore tarsomeres, and 2nd–5th mid and hind tarsomeres dark; tarsal claws reddish brown. Forewings (Fig. 23) pale smoky brown, with 1 yellowish marking at base, 4–5 mostly connected yellowish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with brownish stripes; veins pale yellow except basal anal veins and veins on apical half pale brown. Hindwings almost hyaline; veins pale yellow except pale brown veins at tip. Rs 10- branched, last branch trifurcate; 10–12 crossveins between R1 and Rs; M1+2 5 to 6-branched, M3+4 3 to 4- branched; 1A 3-branched. Abdomen dark with yellowish short setae. Male ninth tergum (Fig. 143) subquadrate, with parabolic anterior margin and truncate posterior margin. Ninth sternum (Fig. 144) broad with curved lateral margin and truncate posterior margin; central portion inflated. Ninth gonostylus (Fig. 145) unguiform and slightly curved dorsomedially. Tenth tergum (Figs. 143–144) short and subcylindrical, ventrally with one well developed tufted tubercle near posteromedial corner; posterolateral corner slightly and acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 146) arched, anteriorly elevated at middle; posterior incision extremely small and V-shaped, forming two triangular processes; lateral lobes digitiform with distal half rather narrowed and apparently incurved. Female eighth sternum (Figs. 147–148) strongly sclerotized, subtriangular in lateral view, posterior margin elevated with moderately developed median incision, forming pair of obtuse processes; ninth segment (Fig. 147) laterally inflated in one pair of large subquadrate sac-like lobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangular upper and one round lower lobe. Materials examined. Holotype %, CHINA: Yunnan, Yunxian, 1100 m, 20.VI.1980, Ming Zhang (CAU). Paratype 1 without genitalia, CHINA: Yunnan, Tengchong (CAU). &, CHINA: Yunnan, Weishan, Weibaos- han, 1800 m, 24.VII.2003, Kunjie Chen (CAU); 2 % 2 &, CHINA: Sichuan, Xichang, Lushan, 2280 m, 5.VIII.1974 (IZCAS).

PROTOHERMES COSTALIS SPECIES-GROUP Zootaxa 1439 © 2007 Magnolia Press · 43 Distribution. China (Sichuan, Yunnan). Remarks. After our examination of the types of this species, we found that the male genitalia of the holotype as well as the appearance are rather different from its original description. However, the features of the four male paratypes differ greatly from the original description. So, based on the four male paratypes of P. yunnanensis and some other material, we established a new species, P. triangulatus, as described above. Re- description of P. yunnanensis was made based on the holotype male and some other materials from Sichuan and Yunnan. Protohermes yunnanensis appears to be closely related to P. costalis in having the same pattern of markings on head, thorax and wings, but the general body coloration in P. yunnanensis is much paler than P. c o st a - lis. In addition, P. yunnanensis is unique in the form of the male ninth sternum with the truncate posterior margin. In P. co s ta l is, the male ninth sternum has a deep trapezoidal posterior incision.

FIGURES 142–148. Protohermes yunnanensis Yang & Yang. 142. Male head and pronotum, dorsal view. 143. Male genitalia, dorsal view. 144. Male genitalia, ventral view. 145. Male ninth gonostylus, caudal view. 146. Male tenth sternum, ventral view. 147. Female genitalia, lateral view. 148. Female eighth sternum, ventral view. Scale lines = 1.0 mm

Acknowledgements

We are grateful to Dr. N. D. Penny (California Academy of Sciences), Dr. S. L. Winterton (California Depart- ment of Food & Agriculture), and Dr. Z. L. Sha (Chinese Academy of Science) for their critical comments after their proof reading on the manuscript. Our thanks also go to Mr. J. Yao, Mr. X. W. Liu, Mrs. W. N. Zhang (Chinese Academy of Science), and Mr. L. Tang (Shanghai Normal University) for providing many speci-

44 · Zootaxa 1439 © 2007 Magnolia Press LIU ET AL. mens from their Insect Collections. We are also much indebted to Mr. F. S. Li (Beijing) and Dr. O. S. Flint, Jr. (Washington, D. C.) for their kind help in many ways. This research was supported by the National Natural Science Foundation of China (30370174, 30225009).

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