Protohermes costalis species-group

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

ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2007 · Magnolia Press

Zootaxa 1439: 1–46 (2007) www.mapress.com/zootaxa/

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] of Biology, Tokyo Metropolitan University, Minamiosawa 1–1, Hachioji, Tokyo 192–0397, Japan.*Corresponding author

Table of contents

Abstract ............................................................................................................................................................................... 2Introduction .........................................................................................................................................................................2Material and methods .......................................................................................................................................................... 2Results .................................................................................................................................................................................7Discussion ........................................................................................................................................................................... 7Systematics........................................................................................................................................................................ 11Genus Protohermes Weele 1907 ....................................................................................................................................... 11

The Protohermes costalis species-group ....................................................................................................................11Key to species of the P. costalis species-group 12

Protohermes acutatus sp. nov. ....................................................................................................................................13Protohermes arunachalensis Ghosh ..........................................................................................................................16Protohermes basiflavus Yang .....................................................................................................................................18Protohermes basimaculatus sp. nov. .........................................................................................................................19Protohermes costalis (Walker) ...................................................................................................................................21Protohermes disjunctus sp. nov. ................................................................................................................................23Protohermes fujianensis Yang & Yang ......................................................................................................................25Protohermes gutianensis Yang & Yang .....................................................................................................................26Protohermes hunanensis Yang & Yang ......................................................................................................................28Protohermes lii sp. nov. .............................................................................................................................................30Protohermes niger Yang & Yang ...............................................................................................................................31Protohermes orientalis sp. nov. .................................................................................................................................33Protohermes similis Yang & Yang .............................................................................................................................35Protohermes sinensis Yang & Yang ...........................................................................................................................36Protohermes stigmosus sp. nov. .................................................................................................................................38Protohermes triangulatus sp. nov. .............................................................................................................................39Protohermes yangi sp. nov. ........................................................................................................................................41Protohermes yunnanensis Yang & Yang ....................................................................................................................43

Acknowledgements ...........................................................................................................................................................44References .........................................................................................................................................................................45

LIU ET AL.2 · Zootaxa 1439 © 2007 Magnolia Press

Abstract

The dobsonfly genus Protohermes is the largest genus in Megaloptera, with the richest species diversity and the widestgeographic distributions. In the present study, the Protohermes costalis species-group is recognised, including 18 spe-cies. Eight species are described herein as new: Protohermes acutatus, Protohermes basimaculatus, Protohermes dis-junctus, Protohermes lii, Protohermes orientalis, Protohermes stigmosus, Protohermes triangulatus and Protohermesyangi 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 describedspecies, which are distributed from far eastern to southeastern Asia. The adult is medium to large sized, usu-ally with yellowish body coloration and several yellowish or whitish markings on wings. The male genitaliashow various modifications and provide useful evidence for specific identification. The male tenth tergumvaries considerably from elongated band-like structures to short subcylindrical processes. The male ninth ster-num is modified to form a subgenital plate, with a posterior incision varying from deep to shallow and fromV-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 mountain-ous and wooded areas. The adults emerge from late spring to early autumn, are usually found near streambanks, and are readily collected at lights. The detailed natural history of Protohermes is well known only inProtohermes 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 asits sister. Together they are a sister lineage to the remaining genera within Corydalinae under Penny’s assump-tion (1993) or sister lineage to the rest of Corydalinae genera, excepting Chloroniella, with the most primitivephylogenetic status under Contreras-Ramos’ (1998) assumption. However, the interspecific phylogeny of theProtohermes species remains poorly studied so far. Recently, Liu & Yang (2005) have split the genus into sev-eral species groups.

Protohermes costalis (Walker) is a common species in Oriental China. It is characterized by widely sepa-rated ocelli and a subcylindrical male tenth tergum bearing a ventral tuft. During this study, more species werefound to be closely related to P. costalis. Thus, in the present paper, the Protohermes costalis species-group isrecognized. All 18 group members, including eight species new to science, are keyed, described, and illus-trated. A preliminary phylogenetic analysis of the P. costalis species group is undertaken based on morpholog-ical data. Geographic distributions are updated, and phylogenetic relationships and biogeography for thespecies group are also discussed.

Material and methods

Taxa studiedSpecimens were obtained from the following collections: the Entomological Museum of China Agricul-

tural University (CAU), Beijing; Hebei University (HBU), Baoding; Institute of Zoology, Chinese Academyof Sciences (IZCAS), Beijing; Kunming Institute of Zoology, Chinese Academy of Sciences (KIZCAS), Kun-

Zootaxa 1439 © 2007 Magnolia Press · 3PROTOHERMES COSTALIS SPECIES-GROUP

ming; the Shanghai Entomological Museum, Chinese Academy of Sciences (SEMCAS), Shanghai; ShanghaiNormal 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 8–10hours. The abdomen was transferred to glycerin for further examination and moved to a microvial with freshglycerin, pinned below the specimen. Habitus photos of adults were obtained using a Nikon Coolpix 4500digital camera. Drawings were made under the light microscope. Terminology generally follows that of Con-treras-Ramos (2004).

Cladistic analysisThe phylogenetic analysis includes 35 mophological characters (Table 1) obtained from all 18 species of

the P. costalis group. The cladogram was rooted using the following two Protohermes species, Protohermesinfectus (McLachlan) and Protohermes xanthodes Navás. Although the phylogenetic relationships among allthe Protohermes species have not been worked out, Protohermes infectus is considered herein to be a speciespossessing relatively more plesiomorphic characters than the other Protohermes species, such as the closelygathered ocelli, the male ninth gonostylus not eveloped by the ninth tergum, and the simple digitiformed maletenth tergum. Protohermes xanthodes seems to have the closest affinity with the P. costalis group in havingsimilar 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 par-simony 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 searchingwas 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 MacCladeversion 4.0 (Maddison & Maddison, 2000). Characters and character states employed in this analysis arelisted below. Character states were scored 0–2 or ? (0 = plesiomorphic state, 1–2 = apomorphic state, ? = stateunknown).

1. Head with post-ocular spine: (0) Present; (1) Absent. The post-ocular spine is present in many generaof Corydalinae. Most Protohermes species have a pair of post-ocular spines, though they are much weakerthan that in Acanthacorydalis, Neoneuromus, etc. In most species of the P. costalis group the post-ocular spineis absent except in P. basimaculatus sp. nov., P. niger Yang & Yang, and P. stigmosus sp. nov. with obtusepost-ocular spines (Figs. 45, 95, 123).

2. Posterior ocelli: (0) Close to each other; (1) Widely separated from each other. In P. xanthodes and allspecies of the P. costalis group, the ocelli are distinctly widely separated, mostly with the distance between theposterior 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 brownwithout 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) Entirelyblack. In most P. costalis group members, there are at least two markings on each side, while there is only onevitta in P. fujianensis Yang & Yang, P. gutianensis Yang & Yang, P. hunanensis Yang & Yang, P. sinensis Yang& Yang, and P. yangi sp. nov., and the pronotum is entirely black in P. niger.

5. Forewing with round markings on apical 1/3: (0) With two radial veins at most through them; (1) Withthree or four radial veins through them. The round marking on forewing is an apomorphic character in manyProtohermes and all Neurhermes species. The normal form of the apical markings is round with only tworadial veins through them. However, in P. fujianensis, P. hunanensis, and P. yangi, the round markings areenlarged, with three or four radial veins through them (Figs. 12, 14, 22).


TAB

LE

1. C

hara

cter

stat

e di

strib

utio

n fo

r the

Pro

tohe

rmes

cos

talis

gro

up u

sed

in th

e cl

adis

tic a

naly

sis.

Cha

ract

ers

12

34

56

7 8

9

1 01

23

45

67

89

2 01

23

45

67

89

3 01

23

45

Taxa

P. in

fect

us

P. x

anth

odes

P. a

cuta

tus

P. a

runa

chal

ensi

s

P. b

asifl

avus

P. b

asim

acul

atus

P. c

osta

lis

P. d

isju

nctu

s

P. fu

jiane

nsis

P. g

utia

nens

is

P. h

unan

ensi

s

P. li

i

P. n

iger

P. o

rien

talis

P. si

mili

s

P. si

nens

is

P. st

igm

osus

P. tr

iang

ulat

us

P. y

angi

P. y

unna

nens

is

0 0 1 1 1 0 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 0 0 1 0 1 1 0 0 0 1 0 0 0 0 1 0 0 1

0 0 0 0 0 0 0 0 1 1 1 0 2 0 0 1 0 0 1 0

0 0 0 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 1 0

1 0 1 1 1 2 1 1 1 1 1 1 0 1 1 0 1 1 1 1

0 0 1 1 1 0 1 1 1 1 1 1 0 1 1 1 0 0 1 1

0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 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

0 1 1 1 1 0 0 1 1 1 1 1 0 1 1 0 0 0 0 1

0 0 1 0 1 0 1 1 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 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0

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

0 0 1 0 1 0 1 0 0 1 0 0 1 0 0 0 0 1 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 1 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 0 1 1 1 1 1 1 0 1 1 0 0 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 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 1 1 1 1 1 1 1 0 1 1 1 1 1 1 1

0 0 1 1 1 0 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 0 0 0 0 0 0 0 0

0 0 1 0 1 0 1 1 0 0 0 1 0 0 0 0 2 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 1 1 1 1 1 1 1 0 1 1 1 0 1 1 1

0 0 1 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 1

0 0 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 1 1 ? 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 0 0 0 0 ? ? 0 0 0 ? 0 0 1


6. Forewing with stripes on costal areas: (0) Absent; (1) Present in most cellules; (2) Present in only basal1–3 cellules. The immaculate forewing costal areas is apomorphic, while there are numerous dark stripes inthe forewing costal cellules in many Protohermes species. However, in some Protohermes species, the darkstripes 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 plesiomor-

phic character in most Protohermes species, while it is three or four branched in many species of the P. costa-lis group, which is considered to be apomorphic.

8. Male ninth sternum, posterior margin: (0) Incised;(1) Truncate. Penny (1993) notes that the posteriorlyincised male ninth sternum is an apomorphic character in Protohermes, but in P. disjunctus and P. yunnanensisYang & Yang, it is almost truncate (Figs. 62, 144).

9. Male ninth sternum, posterior incision: (0) Trapezoidal; (1) V-shaped. In the outgroups and most spe-cies of the P. costalis group, the posterior incision of the male ninth sternum is trapezoidal except for P. basi-maculatus, P. sinensis, 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 trap-ezoidal posterior incision in the present analysis.

10. Male ninth sternum, lateral margin: (0) Straight; (1) Curved. In P. infectus, P. basimaculatus, P. costa-lis, P. niger, P. sinensis, P. stigmosus, P. triangulatus, and P. yangi, the lateral margin of the male ninth ster-num 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. costalis, 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 theother species of the P. costalis group.

12. Male ninth sternum, posterior processes: (0) Widely separated; (1) Close to each other. The posteriorprocess of the male ninth sternum is formed by the posterior incision of the sternum. In P. infectus, P. basifla-vus, P. basimaculatus, P. orientalis sp. nov., P. sinensis, and P. stigmosus, the posterior processes are widelyseparated (Figs. 40, 46, 103, 117, 125), while in P. xanthodes and the other species of the P. costalis group, theprocesses are close to each other.

13. Male ninth sternum, length from base to tip of posterior process: (0) As long as ninth tergum; (1) Aslong as ninth plus tenth tergum. In P. acutatus and P. costalis, the male ninth sternum is long, with its lengthfrom 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 maleninth gonostylus is well developed and visible in dorsal view. In the P. costalis group, the male ninth gonosty-lus 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. niger, and P. yangi, the male ninth gonostylus is strongly curved inwards anddorsad (Figs. 49, 63, 70, 85, 98, 138), while the male ninth gonostylus is slightly curved in the outgroups andin the other species of the P. costalis group.

16. Male tenth tergum: (0) Not cylindrical; (1) Subcylindrical. The short subcylindrical male tenth tergumis synapomorphic to the P. costalis group.

17. Male tenth tergum, ventral process: (0) Feebly developed; (1) Well developed. The ventral process ofthe tenth tergum is an additional process located at the inner margin or ventral surface of the tenth tergum insome Protohermes species that often bears a tuft on the male tenth tergum. In P. acutatus, P. basiflavus, P. cos-talis, P. gutianensis, P. niger, P. triangulatus, and P. yunnanensis, the ventral process is well developed, whilein 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. costa-lis group, there is one small tuft on the male tenth tergum (Fig. 26).


19. Male tenth tergum, position of tuft: (0) At inner margin of tenth tergum; (1) At ventral surface of tenthtergum. In P. xanthodes, the tuft is located at the inner margin of the male tenth tergum, whereas in all the spe-cies 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 maletenth tergum is not incised at tip, while the male tenth tergum has its tip more or less incised in the other spe-cies of the P. costalis group.

21. Male tenth tergum, posterolateral corner: (0) Not produced; (1) Produced. The posterolateral cornerof the male tenth tergum is more or less produced outward in most species of the P. costalis group, while it isnot 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 pro-duced. In P. hunanensis and P. yangi, 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. niger,the male tenth tergum is somewhat flattened, while in the other species of the P. costalis group, the male tenthtergum is not flattened.

24. Male tenth ternum: (0) Slightly narrowed toward apex; (1) Not narrowed. In the outgroups, P. nigerand P. stigmosus, the male tenth tergum is slightly narrowed toward its apex (Figs. 96, 124), while in the otherspecies 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 withthe anterior margin elevated is shared by most species of the P. costalis group except P. niger.

26. Male tenth sternum with posterior margin medially incised: (0) Not incised; (1) Incised. In most spe-cies of the P. costalis group, the male tenth sternum is medially incised on the posterior margin, while in theoutgroups 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, andP. gutianensis, the lateral arm of the male tenth sternum is long (Figs. 28, 42, 57, 79), while in the other spe-cies 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 islocated on the sides of the posterior incision and present in a few species of the P. costalis group. In P. acuta-tus, P. basiflavus, P. costalis, P. disjunctus, P. lii, and P. yunnanensis, the posterior process is distinct but rathersmall (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 ofthe P. costalis 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. costalis 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. gutianen-sis, 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. costalis, P. disjunctus, P. orientalis, P.similis, P. sinensis, and P. yunnanensis.

34. Female ninth segment, sac-like lobes: (0) Absent; (1) Present. The sac-like lobe is laterally inflatedfrom the female ninth abdominal segment, which is present in most species of the P. costalis group. However,the sac-like lobe is absent in the outgroups and in P. basimaculatus. During this study, we found some femalesof P. arunachalensis and P. gutianensis retained an incomplete spermatphore attached to its genitalia, held pri-


marily by the lateral sac-like lobes of the ninth sternum. So, perhaps the sac-like lobes play a more importantrole in mating than the other female genitalic structures, and hence are much more modified and may be usedas 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. similis, P. sinensis, P. yangi, and P. triangulatus, thelateral sac-like lobe of the female ninth segment is feebly developed, with its diameter about 0.5–0.75 mm atmaximum (Figs. 36, 65, 73, 80, 88, 106, 113, 133, 140). In P. acutatus, P. basiflavus, P. costalis, P. similis, andP. 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 unambig-uous characters have been mapped directly on the MP tree. An additional cladogram with >50% bootstrap val-ues is shown as Figure 2. According to the MP tree and the tree with the bootstrap values, the P. costalis groupwas found to be monophyletic. Within the ingroup, P. niger was assigned as the sister taxon with all the othergroup members. Similar to P. niger, three species, P. stigmosus, P. basimacualtus, and P. sinensis, were placedto be the sister taxa with the remaining species below their own nodes. Except for the former four species, theremaining ingroup species formed a monophyletic subgroup and separated into two smaller groups containingP. acutatus, P. basiflavus, P. costalis, P. disjunctus, P. lii, P. orientalis, P. similis, and P. hunanensis, P. yangi, P.fujianensis, and P. gutianensis. However, this subgroup is internally paraphyletic in Figure 2.

Discussion

Monopyly of the P. costalis groupThe monophyly of the P. costalis 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 synapomor-phic characters, the sac-like lobe of the female ninth abdominal segment might be a synapomorphic characterof 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. assa-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 beenworked out. However, the close relationships among the P. assamensis, the P. changninganus, the P. costalis,and the P. xanthodes groups might be preliminarily inferred by the widely separated ocelli, the tuft of the maletenth tergum, both of which are shared by the P. costalis and P. xanthodes groups, and by the dorsally invisiblemale ninth gonostylus, which is shared by the P. assamensis, P. changninganus, and P. costalis groups.

Interspecific relationships of the P. costalis groupThe four ingroup taxa, P. niger, 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 eachother, which implies that these species might be representatives of the remnants from former diverse lineagesor have evolved for a long time.


FIGURE 1. The most parsimonious tree of the P. costalis group with the Bremer’s decay indices indicated at nodes (seetext 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 areplaced above the squares.

Excluding the above four ingroup taxa, the monophyletic group comprising of the other ingroups wassupported by the posterior processes of the male ninth sternum close to each other (char. 12: 1) and acutelyproduced posterolateral corner of the male tenth tergum (char. 21: 1, char. 22: 1) of the male ninth sternum.The subclade, which contains P. similis, P. orientalis, P. disjunctus, P. lii, P. yunnanensis, P. basiflavus, P. cos-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. costalis, and P. acu-tatus was supported by the immaculate vertex (char. 3: 1, CI = 0.333) and the presence of the posterior processon the male tenth sternum (char. 28: 1, CI = 1.000). So, in P. acutatus, the weak vertex marking is consideredto reflect an intermedial state during degenerating of the markings.

Another subclade comprising of P. gutianensis, P. fujianensis, P. hunanensis, and P. yangi was supportedby 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. similis and P. costalis, such as the round lateral margins of the vertex, the indistinctwing 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 abnor-mally darkened in many individuals of P. fujianensis, P. hunanensis, and P. yangi, which may also indicate a


common evolutionary history for these species, even though this character was not used in the cladistic analy-sis.

However, considering the subgroup including 14 group members, the Bremer’s decay values of severalbranches are relatively low and the internal topology is paraphyletic after bootstrapping, which might beexplained 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 aquatichabitats of these corydalid species, perhaps causing a conservative morphology at the species level. Thus, theconservative morphological structures, particularly in their genitalia, might lead to considerable difficulties infinding enough informative characters for cladistic analysis.

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

BiogeographyAll species of the P. costalis 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. basifla-vus, and P. costalis are also distributed in the neighboring countries and P. disjunctus only occurs on Ishigakiand Iriomote Islands of southern Ryukyus, Japan. Considering the Chinese fauna, the species of the P. costalisgroup are distributed in four zoogeographical regions (Zhang, 1998), showing different interspecific distribu-tion patterns. In the South China Region, 13 species reside (P. arunachalensis, P. basiflavus, P. basimaculatus,P. costalis, P. gutianensis, P. hunanensis, P. lii, P. niger, P. orientalis, P. stigmosus, P. triangulatus, P. yangi,and P. yunnanensis), and nine of them (P. arunachalensis, P. basiflavus, P. basimaculatus, P. lii, P. niger, P.stigmosus, P. triangulatus, P. yangi and P. yunnanensis) are endemic. In the Southwest China Region, thereare three species, P. gutianensis, P. similis, and P. yunnanensis, with P. similis 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. costalis (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. similis

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. niger Yang

& Yang (�).

3

4

5


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 spe-cies, P. costalis, P. fujianensis, P. gutianensis, P. orientalis, and P. sinensis, with P. fujianensis and P. sinensisendemic to the subregion. The latter subregion also contains five species, P. acutatus, P. costalis, P. gutianen-sis, P. hunanensis, and P. orientalis. Protohermes acutatus is the single palaearctic representive of the P. cos-talis group in the North China Region.

China, particularly the South China Region, is considered to be the distribution center of the P. costalisgroup since it has the richest species diversity. Previous studies on the zoogeography of Megaloptera showedthat the current Asian Corydalidae fauna may originate from northeastern India or southwestern China duringthe 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 duringthe split of Laurasia and Gondwanaland but not the connection of the India subcontinent to Eurasia accordingto the study on the generic phylogeny of Chauliodinae (Liu & Yang, 2006c). However, the center of origin ofthe Asian Corydalidae is corroborated by several independent studies on the Asian megalopteran fauna (Liu &Yang, 2005, 2006a; Liu et al., 2005).

Considering the P. costalis group, P. niger, P. stigmosus, P. basimaculatus, and P. triangulatus, whichoccupy the relatively basal phylogenetic position, were distributed in Yunnan, which consists of the westernpart of the South China Region and the southern part of the Southwest China Region. So, the center of originof the P. costalis group might also fit with that of the Asian Corydalidae mentioned previously. The earlierspeciation of the P. costalis 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. costalis group is herein recognized from the center of origin to the northernand eastern areas. This northward dispersal may have occurred through the Southwest China Region to theNorth China Region, with P. acutatus as the representive of the northward invasion. Eastward dispersal mayhave occurred through the western parts of the South and North China Region, to the eastern parts of theSouth and North China Region. The occurrence of P. costalis in Taiwan and P. disjunctus in the southernRyukyus imply the eastward distribution track of the P. costalis group. Thus, the close relationships among theAsian mainland, Taiwan, and southern Ryukyus might be inferred from the similar fauna of the P. costalisgroup.

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 betweenthem mostly equal to that between antennal bases. Male ninth gonostylus always enveloped by ninth tergumand invisible in dorsal view. Male tenth tergum short and subcylindrical, ventrally usually produced into atubercle 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 sub-trapezoidal and narrowing toward apical portion. Female ninth abdominal segment mostly inflated laterallyforming a pair of sac-like lobes.


Key to species of the P. costalis species-group

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 ...................... 22. Head with post-ocular spine (Figs. 45, 95, 123)......................................................................................... 3- Head without post-ocular spine .................................................................................................................. 43. Pronotum dark (Fig. 95) ............................................................................................ .P. niger 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. sinensis Yang & Yang5. Vertex without any blackish markings......................................................................................................... 6- Vertex with blackish markings near lateral sides....................................................................................... 106. Male ninth sternum with posterior incision ................................................................................................. 7- Male ninth sternum with posterior margin truncate (Figs. 62, 144) ............................................................ 97. Male ninth sternum with rather shallow posterior incision (Fig. 40) ...............................P. basiflavus Yang- Male ninth sternum with deep posterior incision (Figs. 55, 92) .................................................................. 88. 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 ventralturbercle; 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 ............................................................................ 1611. 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 ............................................................................ 1313. 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............................................................... 1414. Forewing with median yellowish markings mostly connected (Fig. 7); male tenth tergum with posterolat-

eral corner distinctly produced (Fig. 32) ............................................................... P. arunachalensis Ghosh- Forewing with median yellowish markings separated (Fig. 18); male tenth tergum with posterolateral cor-

ner 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) ....................... 1616. 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........................................................... 1717. Mandible and antenna entirely black; lateral lobes of male tenth sternum slightly incurved (Fig. 87) ........

...........................................................................................................................P. hunanensis Yang & Yang


- Mandible yellow with apical half black; antenna black with scape and pedicel yellow; lateral lobes ofmale 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 blackishmarkings on each side; male tenth sternum with well developed V-shaped posterior incision, leaving twoacutely angled, triangular distal processes.

Body length 30–32 mm (male) and 43–45 mm (female); forewing length 41–42 mm (male) and 48–50mm (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 mark-ings 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 andpedicel yellowish brown. Mouthparts yellowish brown; maxillary palpus and labial palpus dark brown, man-dible with distal half black.

Prothorax (Fig. 24) yellowish brown; pronotum with two blackish markings on each side. Meso- and met-athorax pale yellow to yellowish brown, each dorsally with one pair of brownish markings laterally. Thoracicpilosity yellow, much longer on meso- and metathorax. Legs yellow, with dense, yellowish, short setae; tibiaeblack apically; tarsal claws reddish brown. Forewings (Fig. 6) pale brown, with 1 large yellowish marking atbase, 3–4 yellowish markings at middle, and 1 extremely small white spot on apical 1/3; costal cellules withindistinct 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; 9–10crossveins 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 trapezoidalanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 26) broad; posterior margin withdeep 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; postero-lateral 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 twoacutely angled triangular processes; lateral lobes elongated and digitiform, distally slightly incurved. Femaleeighth sternum (Fig. 29–30) strongly sclerotized and subtriangular in lateral view; apical margin distinctlyelevated with arcuate median incision, forming pair of obtuse processes; ninth segment (Fig. 29) laterallyinflated in one pair of large subquadrate sac-like lobes; ninth gonocoxite valvate and membranous with onedigitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one subtriangularupper 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.


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. costalis (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.


FIGURES 16–23. Habitus of the species in the P. costalis group. 16. P. niger Yang & Yang (Holotype), male. 17. P. ori-entalis sp. nov. (Holotype), male. 18. P. similis Yang & Yang (Holotype), male. 19. P. sinensis Yang & Yang (Holotype),male. 20. P. stigmosus sp. nov. (Holotype), male. 21. P. triangulatus sp. nov. (Holotype), male. 22. P. yangi 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. similis in having a similar pattern of mark-ings on the head and prothorax, while it is distinguished by the pattern of markings on the wings and the struc-ture of the male genitalia. In P. acutatus, the yellowish round markings on the apical 1/3 of the forewings areabsent, instead having an extremely small white spot, and the posterior incision of the male tenth sternum isdistinct, forming a pair of acutely angled posterior processes. In P. similis, the round yellowish markings aredistinct on the apical 1/3 of the forewings, and the posterior incision of the male tenth sternum is extremelysmall.


FIGURES 24–30. Protohermes acutatus, sp. nov. 24. Male head and pronotum, dorsal view. 25. Male genitalia, dorsalview. 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; maleninth 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, posterolateralmarkings subtriangular. Occiput laterally with pair of black markings. Compound eyes greyish brown; ocelliyellow, darkly margined medially. Antennae dark, with scapes and pedicel yellow. Mouthparts yellow; mandi-ble with distal half black.

Thorax yellow. Pronotum (Fig. 31) laterally with two pairs of large black markings; posterolateral cornerblack. Meso- and metanota brownish on lateral sides. Thoracic pilosity yellow, much longer on meso- andmetathorax. Legs yellow with dense yellowish short setae; fore and middle tibiae with dorsal surface black,


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 mid-dle, and 1 round yellowish marking on apical 1/3; costal cellules with distinct brown stripes. Hindwings muchpaler than forewings, with basal half almost hyaline, and with 1 round yellowish marking on apical 1/3. Veinspale 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, ventralview. 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, witharched anterior and slightly incised posterior margin. Ninth sternum (Fig. 33) broad with arched lateral mar-gin, posterior margin with trapezoidal incision forming two blunt triangular posterior processes; central por-tion apparently inflated. Ninth gonostylus (Fig. 34) unguiform and slightly curved dorsomedially. Tenthtergum (Figs. 32–33) short subcylindrical, ventrally with one feebly developed tufted tubercle at posterome-dial 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 oneextremely 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) laterallywith a pair of small suboval sac-like lobes; ninth gonocoxite valvate and membranous with one digitiformprocess at tip; tenth tergum short, with posterior margin medially incised, leaving one short subtriangularupper and one semicircular lower lobe.


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 anti-cus (Walker). Although the type specimen was not available for examination, the presence of a subcylindricalmale 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 asP. arunachalensis, agreeing closely with the original description. The redescription of the male and descrip-tion of the female was based on the specimens from western Yunnan. This species seems to be closely relatedto 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 ourspecimens as well as in the other species of the P. costalis 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 ninthsternum 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 eyespale 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- andmetathorax 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, withnarrow 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 yellow-ish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with distinct brownstripes. Hindwings much paler than forewings, respectively with 1 round yellowish marking at middle andapical 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 trapezoi-dal anterior margin and truncate or sightly incised posterior margin. Ninth sternum (Fig. 40) broad witharched lateral margin, posterior margin with rather shallow trapezoidal incision, forming two blunt posteriorprocesses; central portion apparently inflated. Ninth gonostylus (Fig. 41) unguiform and slightly curved dor-somedially, 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 andacutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 42) arched, anteriorly slightly


elevated at middle; posterior incision extremely small and V-shaped, forming two papillated processes; laterallobes digitiform and slightly incurved. Female eighth sternum (Figs. 43–44) strongly sclerotized, subtriangu-lar in lateral view, posterior margin distinctly elevated with distinct median incision, forming pair of obtuseprocesses; ninth segment (Fig. 43) laterally inflated in one pair of large subquadrate sac-like lobes; ninthgonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posteriormargin 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


markings; costal areas of forewing with basal 1–3 cellules marked brown; male tenth tergum laterally flat-tened; 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 mark-ings subtriangular, medial markings suboval. Occiput laterally with pair of black markings. Compound eyesgreyish 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.


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 yel-low, 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 brownmarkings. Hindwings much paler than forewings, with 1 rather indistinct round yellowish marking on apical1/3. Veins pale yellow but slightly darker toward apex; costal crossveins dark. Rs 9 to 10-branched, lastbranch 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 ante-rior 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; cen-tral 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 posteromedialcorner; tip slightly incised and densely setose. Tenth sternum (Fig. 50) arched, anteriorly slightly elevated atmiddle; 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 trun-cate; ninth segment (Fig. 51) without sac-like lobes; ninth gonocoxite valvate and membranous with one dig-itiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one short digitiformupper 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 femaleninth 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 sternumwith deep trapezoidal posterior incision, leaving two acute-angled triangular posterior processes.


Head (Fig. 53) yellowish brown, without post-ocular spine; vertex without any markings. Compound eyespale yellowish brown; ocelli yellow, darkly margined medially. Antennae black, with scapes and pedicel yel-low. 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 metanotaeach 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; tarsal claws reddishbrown. Forewings (Fig. 10) pale smoky brown, with 1 large yellowish marking at base, 3–4 mostly connectedyellowish markings at middle and 1 round yellowish marking on apical 1/3; costal cellules with distinct brownstripes. Hindwings much paler than forewings, with 1 round yellowish marking on apical 1/3. Veins yellowishbrown except veins in yellowish markings yellow. Rs 10-branched, last branch bifurcate or trifurcate; 11crossveins 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 ante-rior and posterior margin. Ninth sternum (Fig. 55) broad with nearly straight lateral margin, posterior marginwith deep trapezoidal incision, forming two acutely angled triangular posterior processes; central portioninflated. 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; postero-lateral corner acutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 57) arched, anteri-orly slightly elevated at middle; posterior incision extremely small and V-shaped, forming two papillatedprocesses; 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 shallowmedian incision, forming pair of obtuse processes; ninth segment (Fig. 58) laterally inflated in one pair oflarge 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 subquad-rate lower lobe.

FIGURES 53–59. Protohermes costalis (Walker). 53. Male head and pronotum, dorsal view. 54. Male genitalia, dorsalview. 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); &,


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, 900m, 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, 650m, 27.VI.1991, Longlong Yang (IZCAS); 2 &, Fujian, Jiangle, Longqishan, 700 m, 1–6.VIII.1991, ShimeiSong (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. costalis is considered to be a valid species and not asynonym of Hermes anticus Walker. For comparison of P. costalis with other related species, see the remarksunder 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 eventruncate; male tenth tergum acutely produced on posterolateral portion, ventrally with feebly developed tuftedtubercle; female abdominal segment with pair of small lateral lobes.


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 pairmuch larger. Meso- and metathorax pale yellow. Thoracic pilosity pale yellow, much longer on meso- andmetathorax. Legs pale yellow with short, dense, yellowish setae; foreleg with wide apex of tibiae dark, tarsidark, tarsal claws reddish brown. Forewings (Fig. 11) slightly grayish brown, with 1 large subtriangular yel-lowish marking at base, 1 large round and several small discrete yellowish markings at middle, and 1 roundyellowish marking on apical 1/3; costal cellules with distinct brownish stripes. Hindwings slightly grayishbrown except basal half hyaline, respectively with 1 round yellowish marking at middle and apical 1/3. Veinsyellowish brown, much darker on apical half, but much paler in yellowish markings. Rs 8 to 10-branched, last


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 mar-gins 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 feeblydeveloped 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 pro-cesses; 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 ofroundly 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 ventrallobe.

FIGURES 60–66. Protohermes disjunctus, sp. nov. 60. Male head and pronotum, dorsal view. 61. Male genitalia, dorsalview. 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, FumioHayashi (CAU). Others material: JAPAN: 5 % 2 &, Iriomote Island, Takana-gawa, emerged from reared larvaein 1987, 1988, Fumio Hayashi (HC); 4 % 3 &, Ishigaki Island, Nagura-gawa, emerged from reared larvae in1988, Fumio Hayashi (HC).


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. cos-

talis during the separation of the Iriomote Island from Taiwan. However, considering the genitalia, P. disjunc-tus appears to be closely related to P. yunnanensis in having a similar male ninth sternum, but it can be easilyseparated from P. yunnanensis by the male tenth tergum with the feebly developed tufted tubercle and thesmall female sac-like lobes. In P. yunnanensis, the male tenth tergum is ventrally produced into one stronglydeveloped 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 lateralmargins; forewings pale brown with several whitish markings; male ninth sternum with trapezoidal posteriorincision; male tenth sternum with lateral lobes distinctly incurved.


Head (Fig. 67) yellowish brown without post-ocular spine, dorsally with one pair of brown markings onlateral sides. Compound eyes brown; ocelli pale yellow, darkly margined medially. Antennae black withscapes and pedicel pale yellow. Mouthparts yellow; mandible with distal half black. Occiput yellowish brownwith 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- andmetathorax. Legs dark with dense yellowish short setae; coxae, trochanters and basal tibiae pale yellow; tarsalclaws reddish brown. Forewings (Fig. 12) pale brown, with 1 round whitish marking at base, 2 round whitishmarkings at middle and 1 distinctly enlarged, whitish round marking on apical 1/3; costal cellules with distinctbrownish stripes. Hindwings pale brown except basal half subhyaline, respectively with 1 round whitishmarking at base, middle and apical 1/3. Veins pale brown, except veins in whitish markings pale yellow. Rs 9to 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 ante-

rior 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; cen-tral 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; posterolat-eral 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 digiti-form with distal half incurved. Female eighth sternum (Figs. 73–74) strongly sclerotized and subtriangular inlateral view, with truncate posterior margin; ninth segment (Fig. 73) laterally inflated in one pair of small ovalsac-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,


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 yel-low, male tenth sternum with lateral lobes incurved in apical half. By contrast, P. hunanensis has the mandibleand 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, ventralview. 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 narrowblackish vittae medially on vertex; pronotum with one pair of blackish vittae on lateral margins; wings pale


smoky brown with several indistinct yellowish spots except for apical regions without spots; male ninth ster-num with suboblong posterior incision.


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 met-athorax yellow, each dorsally with one pair of brownish spots on anterolateral corners. Thoracic pilosity yel-low, much longer on meso- and metathorax. Legs black with dense, pale yellow, short setae; coxae andtrochanters pale yellow; femora brown; tarsal claws reddish brown. Forewings (Fig. 13) slightly smokybrown, with 1 small yellowish marking at base, 2–4 yellowish markings along cubital veins at middle; costalcellules with distinct brownish stripes. Hindwings much paler than forewings with hyaline basal half. Veinspale brown, except pale yellow veins in yellowish markings and base of hindwings. Rs 9–10 branched, lastbranch 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 ante-

rior 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; cen-tral 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 cor-ner; posterolateral corner distinctly and acutely produced; tip slightly incised and densely setose. Tenth ster-num (Fig. 79) arched, anteriorly with one trapezoidal process at middle and posteriorly with one smalltriangular 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 withone digitiform process at tip; tenth tergum short, with posterior margin medially incised, leaving one long dig-itiform 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, XuezhongZhang (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, 800m, 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 &


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 ofslender blackish vittae and the male ninth sternum with deep suboblong apical incision, both of which areunique in the P. costalis 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, ventralview. 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 onepair of blackish vittae on lateral margins; wings dark with distinct whitish markings.


Head (Fig. 82) yellow without post-ocular spine, dorsally entirely black near lateral margins. Compound


eyes brown; ocelli yellow, darkly margined medially. Antennae black. Mouthparts mostly yellow with blackmandible. 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 met-athorax brown, each dorsally with one pair of blackish spots on lateral sides. Thoracic pilosity yellow, muchlonger 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 mark-ings at middle, and 1 distinctly enlarged round whitish marking on apical 1/3; costal cellules with distinctbrownish stripes. Hindwings dark except basal half hyaline, respectively with 1 large round whitish markingat middle and apical 1/3. Veins dark brown, except pale yellow veins in whitish markings and base of hind-wings. 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, ventralview. 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 ante-rior margin and shallowly incised posterior margin. Ninth sternum (Fig. 84) broad; posterior margin withextremely shallow trapezoidal incision, forming two broad roundly produced processes; central portioninflated. 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


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 andslightly 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-likelobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, withposterior 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, WenzhuLi (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. costalis 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. fujianen-sis 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; maleninth sternum with wide, shallow posterior incision; male tenth tergum with feebly developed ventral tuftedtubercle.

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; man-dible distally dark.

Prothorax yellow; pronotum (Fig. 90) with two pairs of widely separated small blackish markings on eachside. 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; tarsalclaws reddish brown. Forewings (Fig. 15) pale smoky brown, with 1 large yellowish marking at base, 3mostly 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 thanforewings, respectively with 1 round yellowish marking at middle and apical 1/3. Veins brown except veins inyellowish markings and basal half of hindwings pale yellow. Rs 9 to 10-branched, last branch bifurcate; 10–13crossveins 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 ante-rior margin and sightly incised posterior margin. Ninth sternum (Fig. 92) broad with lateral margin slightlycurved near anterior 1/4, posterior margin with broad, shallow incision, forming two acute-angled posteriorprocesses; 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 nearposteromedial corner; posterolateral corner slightly and acutely produced; tip slightly incised and densely set-ose. Tenth sternum (Fig. 94) slightly arched, anteriorly slightly elevated at middle, posterior margin withextremely small V-shaped median incision and a pair of small denticles; lateral lobes digitiform, with apexstrongly 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,


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 tocollect 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 turber-cle of the male tenth tergum, and the male tenth sternum with the posterior margin bearing a pair of small den-ticles. In P. basiflavus, the body size and the prothoracic markings are much larger, the ventral tufted turbercleof the male tenth tergum is well developed, and the male tenth sternum posteriorly bears a pair of distinctlyproduced 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. Scalelines = 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 indistinctyellowish 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 ocellusslightly transverse; posterior pair widely apart, but distance between them slightly shorter than that between


antennal bases. Antennae black with scapes and pedicel yellowish brown. Mouthparts yellow; mandible dis-tally black.

Prothorax (Fig. 95) dark; pronotum with yellowish brown anterior margin, and with one slender yellowbrown spot near each anterolateral corner and one long hook-shaped yellow brown marking following theformer 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 mid-dle and 1 round whitish marking on apical 1/3; costal area brown. Hindwings grayish brown, with 1 indistinctround whitish marking at base, 1 indistinct round whitish marking at middle and 1 distinct round whitishmarking on apical 1/3. Veins dark, except pale yellow on whitish markings. Rs 7 to 8-branched, last branchbifurcate; 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, dor-sal view. 97. Male ninth sternum, ventral view. 98. Male ninth gonostylus, caudal view. 99. Male tenth tergum, ventralview. 100. Male tenth sternum, ventral view. Scale lines = 1.0 mm.

Abdomen brown with yellowish short setae. Male ninth tergum (Fig. 96) subquadrate, with trapezoidalanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 97) broad, posterior margin withshallow trapezoidal incision, forming two broad acute-angled triangular processes; central portion feeblyinflated. 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; postero-lateral corner not produced; tip slightly incised and densely setose. Tenth sternum (Fig. 100) arched with ante-


rior margin feebly elevated; posterior incision triangular and extremely small; lateral lobes thick digitiformand 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, DazhiDong (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, DazhiDong (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. costalis group by the distinct black-

ish pronotum.

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

Diagnosis: Head with three blackish markings on each side of vetex; antenna entirely black; pronotum withtwo blackish markings on each side; legs almost dark; male ninth sternum with extremely shallow trapezoidalposterior incision.


Head (Fig. 101) yellowish brown, without post-ocular spine; vertex with three blackish markings on eachside, anterior marking large and subquadrate, posterolateral marking cuneiform, posteromedial marking punc-tuate. Compound eyes brown; ocelli yellowish brown, darkly margined medially. Antennae black. Mouthpartsyellowish 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- andmetathorax dark, dorsally with each center pale yellow. Thoracic pilosity yellow, much longer on meso- andmeta thorax. Legs dark with dense yellowish short setae; fore and mid legs with trochanters and dorsal surfaceof femora black; tarsal claws reddish brown. Forewings (Fig. 17) pale smoky brown, with 1 yellowish mark-ing at base, 3–4 yellowish markings at middle and 1 round yellowish markings on apical 1/3; costal celluleswith brownish stripes; veins dark except veins in yellowish markings pale yellow. Hindwings much paler thanforewings, respectively with 1 yellowish marking at middle and apical 1/3; veins dark, except veins in yellow-ish 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 anteriormargin and truncate posterior margin. Ninth sternum (Fig. 103) subquadrate, posterior margin with extremelyshallow incision, leaving a pair of small widely separated subtriangular posterior processes; central portioninflated. 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; lat-eral lobes elongated and digitiform, distally incurved. Female eighth sternum (Figs. 106–107) strongly sclero-tized, subtriangular in lateral view, and subtrapezoidal in ventral view, posterior margin slightly incised,


forming pair of obtuse processes; ninth segment (Fig. 106) laterally inflated in one pair of small sac-likelobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, withposterior 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. acuta-tus, 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 (Figs. 18, 108–114)

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 markingson each side; forewings with distinct brownish stripes on costal cellules; male ninth sternum with trapezoidalposterior incision; male tenth sternum with extremely small V-shaped posterior incision and short digitiformlateral lobes.


Head (Fig. 108) pale yellowish brown to reddish brown without post-ocular spine; vertex with one largeblackish marking on each side, occasionally separated into three markings. Compound eyes brown; ocelli yel-low, 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 eachside. 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 yellowishshort 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 1round yellowish marking at middle and apical 1/3. Veins pale brown, except costal crossveins pale yellow andveins 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 trapezoidalanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 110) broad with arched lateralmargins; 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 ter-gum (Figs. 108–109) short and subcylindrical, ventrally with one feebly developed tufted tubercle at postero-medial 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 digiti-form and slightly incurved. Female eighth sternum (Fig. 113–114) strongly sclerotized and subtrapezoidal inlateral view, posterior margin apparently elevated with shallow median incision, forming pair of obtuse pro-cesses; ninth segment (Fig. 113) laterally inflated one pair of small suboval sac-like lobes; ninth gonocoxitevalvate and membranous with one digitiform process at tip; tenth tergum short, with posterior margin medi-ally incised, leaving one thick digitiform upper and one round lower lobe.

Material examined. Holotype %, CHINA: Sichuan, Ya-an, VIII.1984 (CAU). Allotype &, CHINA: samedata 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-


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 speciesis distinguished from P. acutatus by having the forewings with distinct brown stripes on the costal cellules andround yellowish markings at the apical 1/3, while in P. acutatus the forewings are much paler, with indistinctstripes on the costal cellules and the apical portions of the forewings. Protohermes sichuanensis was estab-lished by Yang and Yang (1997) according to one male and one female specimen. After examination to thetype series of P. sichuanensis, no distinct differences from P. similis were found. Contrary to what was statedin the original description, P. sichuanensis has the trapezoidal apical incision on the male ninth sternum,which is the same as P. similis. 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, ventralview. 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).


Diagnosis. Head with one transverse dark marking anterior to ocellar triangle and three blackish markings oneach lateral side of vertex; pronotum with one dark vitta on each lateral side; forewing costal region entirelyhyaline; 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 anteriorto 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 yellow-ish brown; mandible with distal half black.

FIGURES 115–122. Protohermes sinensis Yang & Yang. 115. Male head and pronotum, dorsal view. 116. Male genita-lia, 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 metanotaeach with one pair of blackish markings laterally. Thoracic pilosity yellow, much longer on meso- and met-athorax. Legs dark with dense yellowish short setae; coxae, trochanters and basal tibiae yellow, sometimestibiae entirely brown; tarsal claws reddish brown. Forewings (Fig. 19) pale smoky brown, with 1 yellowishmarking 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 roundyellowish marking at middle and apical 1/3; veins brown, except veins in yellowish markings and basal longi-


tudinal veins yellow. Rs 9 to 11-branched, last branch bifurcate or trifurcate; 9–12 crossveins between R1 and

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 trapezoidalanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 117) subquadrate with straightlateral margin, apical margin with shallow V-shaped incision, forming two small acutely angled apical pro-cesses; 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 pos-teromedial 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; lat-eral lobes digitiform and slightly incurved. Female eighth sternum (Figs. 121–122) strongly sclerotized andsubtrapezoidal in lateral view, posterior margin elevated with shallow median incision, forming pair of obtuseprocesses; ninth segment (Fig. 121) laterally inflated as a pair of small suboval sac-like lobes; ninth gonocox-ite valvate and membranous with a digitiform process on tip; tenth tergum short, with posterior margin medi-ally 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: samedata 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 sepa-

rated from P. xanthodes by the subcylindrical male tenth tergum and the presence of the sac-like lobes in thefemale genitalia. The male ninth sternum of P. sinensis is distinguished from the other species of the P. costa-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 cornerand three pairs of blackish markings at posterolateral corner; male ninth sternum has a wide U-shaped poste-rior incision, leaving two short acutely angled digitiform processes; tenth sternum with posterior incisionstrongly developed, leaving two large papillate processes with slender digitiformed tip, lateral lobes short anddigitiform, 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 andthree 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; tarsalclaws pale reddish brown. Forewings (Fig. 20) pale grayish brown except anal region entirely hyaline, with 1subtriangular yellowish marking near base,1 broad and 1 narrow yellowish markings at middle, 1 round and 1narrow yellowish markings on apical 1/3 and several extremely small yellowish markings on crossveins atapical portion; costal cellules with distinct grayish brown stripes. Hindwings entirely hyaline. Veins pale yel-low except veins in dark regions of forewings and apical portion of hindwings pale brown. Rs 8-branched, last


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 widerposteriorly, 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, form-ing 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 fee-bly 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 medianpapillate processes, lateral lobes slender digitiform; lateral arms short and digitiform, proximally inflated andstrongly 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. costalis group based on

the ten small dark, lateral spots on the pronotum, by the male ninth sternum with a wide, U-shaped posteriorincision, and by the male tenth sternum with the abnormal posterior lobes. However, the more complicatedwing patterns somewhat show the close affinity between P. stigmosus 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, ventralview. 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 blackishmarkings on each side; male ninth sternum with deep V-shaped posterior incision; tenth tergum with an


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 lateralsides.

Thorax yellow. Pronotum (Fig. 128) with two small blackish markings on each side. Meso- and metanotaeach with one pair of brownish markings on lateral sides. Thoracic pilosity yellow, much longer on meso- andmeta thorax. Legs yellow with dense yellowish short setae; tibiae and tarsi mostly dark, and sometimes only2nd–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 onapical 1/3; costal cellules with distinct brownish stripes. Hindwings mostly hyaline, respectively with 1 indis-tinct 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, ventralview. 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 archedanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 130) broad with nearly straightlateral margin; posterior margin with deep V-shaped incision; central portion inflated. Ninth gonostylus (Fig.


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 andacutely produced; tip slightly incised and densely setose. Tenth sternum (Fig. 132) arched, anteriorly slightlyelevated at middle; posterior incision V-shaped and moderately developed; lateral lobes digitiform andslightly incurved. Female eighth sternum (Figs. 133–134) strongly sclerotized and subtrapezoidal in lateralview, with truncate posterior margin; ninth segment (Fig. 133) laterally inflated in one pair of small subovallobes; ninth gonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, withposterior 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, GuoxiuLi (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 themale 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 hasa 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 grayishbrown with distinct whitish markings; male ninth sternum with deep V-shaped posterior incision; male tenthtergum strongly and roundly produced distolaterally.


Head (Fig. 135) yellowish brown without post-ocular spine; vertex with two blackish markings on eachside, 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 palpusbrown, mandible with distal half dark. Occiput yellowish brown with one pair of blackish markings on lateralsides.

Prothorax (Fig. 135) yellowish brown; pronotum with one blackish vitta on each side, vittae sometimesnarrowed 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 yellowishshort setae; wide apical portion of tibiae and tarsi black; tarsal claws reddish brown. Forewings (Fig. 22) gray-ish brown, with 1 subtriangular whitish marking at base, 2 large and 5–6 small whitish markings at middle and1 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 yel-lowish 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.


FIGURES 135–141. Protohermes yangi, sp. nov. 135. Male head and pronotum, dorsal view. 136. Male genitalia, dorsalview. 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 trapezoidalanterior margin and shallowly incised posterior margin. Ninth sternum (Fig. 137) broad with straight lateralmargin; posterior margin with deep V-shaped incision, forming two large triangular processes; central portioninflated. 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. Tenthsternum (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 sclero-tized and subtriangular in lateral view, posterior margin elevated without incision, forming a pair of triangularprocesses; ninth segment (Fig. 140) laterally inflated forming a pair of small suboval sac-like lobes; ninthgonocoxite valvate and membranous with one digitiform process at tip; tenth tergum short, with posteriormargin 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 passedaway in 2006, for his great contributions to the systematics of the Chinese Megaloptera.


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, theposterior incision of the male ninth sternum is trapezoidal in both P. hunanensis and P. fujianensis. The newspecies also somewhat resembles P. triangulatus in having the similar V-shaped posterior incision of the maleninth 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 sternumwith truncate posterior margin.


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 oryellowish brown; mandible with distal half black.

Thorax yellow. Pronotum (Fig. 142) with two narrow blackish markings on each side. Meso- and metan-ota each with posterolateral corners pale brown. Thoracic pilosity yellow, much longer on meso- and metatho-rax. Legs pale yellow with dense yellowish short setae; apical 1/3 of fore tibiae, fore tarsomeres, and 2nd–5thmid and hind tarsomeres dark; tarsal claws reddish brown. Forewings (Fig. 23) pale smoky brown, with 1 yel-lowish marking at base, 4–5 mostly connected yellowish markings at middle and 1 round yellowish markingon apical 1/3; costal cellules with brownish stripes; veins pale yellow except basal anal veins and veins on api-cal 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 ante-

rior margin and truncate posterior margin. Ninth sternum (Fig. 144) broad with curved lateral margin andtruncate posterior margin; central portion inflated. Ninth gonostylus (Fig. 145) unguiform and slightly curveddorsomedially. Tenth tergum (Figs. 143–144) short and subcylindrical, ventrally with one well developedtufted tubercle near posteromedial corner; posterolateral corner slightly and acutely produced; tip slightlyincised and densely setose. Tenth sternum (Fig. 146) arched, anteriorly elevated at middle; posterior incisionextremely small and V-shaped, forming two triangular processes; lateral lobes digitiform with distal halfrather narrowed and apparently incurved. Female eighth sternum (Figs. 147–148) strongly sclerotized, subtri-angular in lateral view, posterior margin elevated with moderately developed median incision, forming pair ofobtuse 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 poste-rior 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).


Distribution. China (Sichuan, Yunnan).Remarks. After our examination of the types of this species, we found that the male genitalia of the holo-

type as well as the appearance are rather different from its original description. However, the features of thefour 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 Sichuanand Yunnan.

Protohermes yunnanensis appears to be closely related to P. costalis in having the same pattern of mark-ings on head, thorax and wings, but the general body coloration in P. yunnanensis is much paler than P. costa-lis. In addition, P. yunnanensis is unique in the form of the male ninth sternum with the truncate posteriormargin. In P. costalis, 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. Malegenitalia, dorsal view. 144. Male genitalia, ventral view. 145. Male ninth gonostylus, caudal view. 146. Male tenth ster-num, 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 commentsafter 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-


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 NaturalScience Foundation of China (30370174, 30225009).

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Protohermes costalis species-group