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The Genus Neochauliodes Van Der Weele (Megaloptera: Corydalidae) from

Indochina, with Description of Three New Species

Article in Annales Zoologici · April 2010

DOI: 10.3161/000345410X499597

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 A N N A L E S Z O O L O G I C I (Warszawa), 2010, 60(1): 109-124

THE GENUS NEOCHAULIODES VAN DER WEELE

(MEGALOPTERA: CORYDALIDAE) FROM INDOCHINA,
WITH DESCRIPTION OF THREE NEW SPECIES

XINGYUE LIU1, 2, FUMIO HAYASHI2 and DING YANG1

1
Department of Entomology, China Agricultural University, Beijing 100193, China;
e-mail: liu_xingyue@yahoo.com.cn
2
Department of Biology, Tokyo Metropolitan University, Minamiosawa 1-1,
Hachioji, Tokyo 192-0397, Japan

Abstract.— A faunistic review of the fishfly genus Neochauliodes van der Weele from
Indochina is given, with 11 species recorded. Amongst them, three species are described as
new to science: Neochauliodes bachmanus sp. nov., Neochauliodes confusus sp. nov.,
and Neochauliodes tamdaoensis sp. nov. Neochauliodes orientalis Yang et Yang, 1992
and Neochauliodes yunnanensis Navás, 1930 are synonymized with Neochauliodes
tonkinensis (van der Weele, 1907).

Key words.— Corydalidae, Chauliodinae, Neochauliodes, new species, Indochina.

INTRODUCTION poorly known with 19 species and subspecies, most of

The fishfly genus Neochauliodes van der Weele,
1909 which includes more than 40 species is the largest
genus of the subfamily Chauliodinae, and is widely dis-
tributed in Oriental Asia, with a few records in eastern
Palaearctic Asia. The genus is characterized by the
antenna with distinct sexual dimorphism, the wings
usually with transversely band-like markings medial-
ly, the straightly directed 1A and 2A, the male
tenth sternum with length equal to the ninth plus tenth
terga.
The taxonomy of the genus Neochauliodes is the
result of work mainly by van der Weele (1905, 1906,
1907, 1909, 1910), Lestage (1927), and Kimmins (1954)
in the first half of last century. However, after that
many taxonomic problems of this genus remain and
faunistic studies are lacking. Recently, the east Asian
fauna of Neochauliodes was intensively studied by Liu
and Yang (2005a, b, 2006), Liu et al. (2007), and Liu
(2008), with 26 species recorded from China, Japan,
and Korean Peninsula. Nevertheless, the fauna of
Neochauliodes in south and southeast Asia are still
which lack revision or redescription since the early
original description.
Indochina, which is composed of Myanmar, Thai-
land, Laos, Vietnam, and Cambodia, is a major compo-
nent of southeast Asia, northwardly connecting to the
mainland China and the Indian subcontinent, and
southwardly extending as a peninsula. Due to the trop-
ical and subtropical rainforest habitat, Indochina is
one of the areas with extraordinarily rich biodiversity
Lestage (1927) made a revision of Megaloptera from
Indochina, in which six species and subspecies of
Neochauliodes were recorded, namely Neochau-
liodes bowringi (Mclachlan), Neochauliodes sim-
plex (Walker), Neochauliodes sinensis sinensis van
der Weele, Neochauliodes sinensis occidentalis van
der Weele, Neochauliodes sinensis meridionalis
van der Weele, and Neochauliodes tonkinensis
(van der Weele). No other revisionary work on
Neochauliodes from Indochina has been made since,
but four species, Neochauliodes umbratus Kimmins,
Neochauliodes moriutii Asahina, Neochaulio-
des koreanus van der Weele, and Neochauliodes

PL ISSN 0003-4541 © Fundacja Natura optima dux

doi: 10.3161/000345410X499597
110 X. LIU, F. HAYASHI and D. YANG
punctatolosus Liu et Yang, were subsequently record-
ed in Indochina (Kimmins 1954, Asahina 1988, Liu et
al. 2007). However, due to the changes of the taxonom-
ic status of many Neochauliodes species and several
misidentifications, the regional fauna of this genus
needs further clarification and more species await
description. We have borrowed Megaloptera from col-
lections worldwide, and have made a research trip to
several European museums. Thus many Neochau-
liodes specimens from Indochina, are available for us
to produce a faunistic revision of this genus from this
area. In the present, 11 species of Neochauliodes are
recorded in Indochina, three of which are described
as new to science. Four new country records are
herein recognized. Additionally, Neochauliodes ori-
entalis Yang et Yang and Neochauliodes yunnan-
ensis Navás are treated as the junior synonyms
of Neochauliodes tonkinensis (van der Weele).
A key to the Neochauliodes species from Indochina is
presented.
MATERIAL AND METHODS
The specimens for the present study are deposited
in several collections worldwide, which are listed as
follows with the suggested acronyms.
BPBM – Bishop Museum, Honolulu;
CAU – Entomological Museum, China Agricultural
University, Beijing;
EUM – Ehime University, Matsuyama;
HBU – Hebei University, Baoding;
HC – Fumio Hayashi, personal collection, Tokyo;
HNHM – Hungarian Natural History Museum, Buda-
pest;
IZCAS – Institute of Zoology, China Academy of Sci-
ence, Beijing;
MFN – Museum für Naturkunde, Berlin;
MNHN – Muséum National d’Histoire Naturelle, Paris;
MZBS – Museu de Zoologia, Ajuntament de Barcelo-
na, Barcelona;
NHM – Natural History Museum, London;
NSMT – National Science Museum, Tokyo;
RMNH – Nationaal Natuurhistorisch Museum, Leiden;
SYSU – Sun Yat-sen University, Guangzhou.
The label data of the primary types and the speci-
mens collected before 1940s with old labels are
presented within quotation marks “ ”. Added infor-
mation that expands or augments the often cryptic
text and provides the geographic coordination in the
labels is always placed in square brackets [ ].
Genitalic preparations were made by clearing the
apex of the abdomen in a cold, saturated KOH solution
for 8–10 h. After rinsing the KOH with acetic acid and
water, the apex of the abdomen was transferred to
glycerin for further dissection and examination. After
examination it was moved to fresh glycerin and stored
in a microvial pinned below the specimen. The termi-
nology of the genitalia follows that of Contreras-Ramos
(2004).
TAXONOMY
Genus Neochauliodes van der Weele
Neochauliodes van der Weele 1909: 259. Type species: Chauliodes
sinensis Walker 1853: 199, original designation.
Key to species of the genus Neochauliodes
from Indochina
1. Posterior second and third veins of Rs on forewing
strongly curved posteriad . . . . . [sinensis group] 2
–. All branches of Rs on forewing smoothly directed
and nearly straight, without distinct curve . . . . . . 6
2. Pronotum brownish . . . . bicuspidatus Liu et Yang
–. Pronotum orange . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Head blackish brown . . . . koreanus van der Weele
–. Head orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Male tenth sternum in lateral view with posterior
half inflated (Liu and Yang 2006: Fig. 25); female
ninth gonocoxite subquadrate (Liu and Yang 2006:
Fig. 28) . . . . . . . . . . . . . punctatolosus Liu et Yang
–. Male tenth sternum in lateral view with posterior
half gradually narrowed (Fig. 16); female ninth
gonocoxite foliate (Fig. 19) . . . . . . . . . . . . . . . . . . . 5
5. Forewings densely dotted by numerous small spots,
medially with a narrow transverse band (Fig. 4);
male tenth sternum in lateral view feebly nar-
rowed posteriad, with a short and robust apex (Fig.
16) . . . . . . . . . . . . . . . . . . . . . . . . . . confusus sp. nov.
–. Forewings sparsely dotted by a few broad mark-
ings, medially with a wide transverse band (Fig. 6);
male tenth sternum in laterally view strongly nar-
rowed posteriad, with a long and slender apex (Liu
and Yang 2005a: Fig. 32) . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . meridionalis van der Weele
6. Forewings proximally with numerous small, brown,
rounded spots on costal area (Fig. 3) [bowringi
group] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
–. Forewings proximally with costal area immaculate
or tinged by a few broad and irregularly shaped
markings (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7. Hindwings medially with a distinct broad trans-
verse band (Fig. 3); male tenth tergum in lateral
view suboval (Liu and Yang 2005a: Fig. 14); female
ninth gonocoxite not produced posteriad (Liu and
Yang 2005a: Fig. 17) . . . . . . bowringi (McLachlan)
–. Hindwings medially without distinct transverse
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA
band (Figs 7, 10); male tenth tergum in lateral view
subquadrate or subtrapezoidal (Liu and Yang 2006:
Figs 14, 17); female ninth gonocoxite somewhat pro-
duced posteriad (Liu and Yang 2006: Fig. 20) . . . . 8
8. Brownish markings on distal half of forewings
indistinct or absent; median plate of male tenth
sternum suboblong in ventral view and in lateral
view without inflation medially (Liu and Yang 2006:
Figs 14–15) . . . . . . . . . . . . . . . . . . moriutii Asahina
–. Brownish markings on distal half of forewings dis-
tinct; median plate of male tenth sternum attenuate
in ventral view and in lateral view with median
inflation (Liu and Yang 2006: Figs 17–18) . . . . . . . .
. . . . . . . . . . . . . . . . . . tonkinensis (van der Weele)
9. Both forewings and hindwings medially with broad
brownish transverse band (Fig. 11); male tenth ster-
num laterally with a pair of additional processes
subdistad on median plate (Liu and Yang 2005a:
Fig. 40) . . . . . . . . . . . . . . . . . . . umbratus Kimmins
–. Forewings and hindwings medially with rather nar-
row and separated transverse band or without
transverse band; male tenth sternum without addi-
tional process on median plate . . . . . . . . . . . . . . . 10
10. Hindwings medially with a short transverse band
(Fig. 9); median plate of male tenth sternum laterally
feebly elevated (Fig. 20) . . . . . tamdaoensis sp. nov.
–. Hindwings medially without transverse band (Fig.
1); median plate of male tenth sternum laterally
strongly and broadly elevated (Fig. 12) . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . bachmanus sp. nov.
Neochauliodes bachmanus sp. nov.
(Figs 1, 12–15, 25)
Type locality. Vietnam (Bach Ma).
Etymology. The specific epithet “bachmanus”
refers to the type locality of the new species.
Description. Male. Body length 26–28 mm; fore-
wing length 37–41 mm, hindwing length 32–36 mm.
Head yellowish brown, slightly tinged with pale
brown on vertex laterally. Compound eyes brown; ocel-
li yellow, medially margined black. Antenna black.
Mouthparts yellowish brown; maxillary and labial palpi
with distal two segments blackish brown.
Prothorax pale brown, pronotum with lateral mar-
gins much darker; meso- and metanota blackish brown.
Legs yellow with short, dense, yellowish setae; femora
with apices blackish brown, tibiae and tarsi blackish
brown except for bases of hind tibiae yellowish brown,
tarsal claws reddish brown. Wings hyaline, slightly
pale brown, with quite a few brownish markings;
pterostigmatical area short, whitish. Forewing slightly
darker on distal half, with distinct blackish brown nyg-
mata; costal area with a dark brown stripe proximally
111
and a much shorter dark brown spot distally to
pterostigmatic area; a series of dark brown small spots
longitudinally dotted along Sc and R1, and another row
of pale brown small spots dotted along stem of Cu1; sev-
eral indistinct brownish spots presented medially and
subdistally, generally forming two narrow transverse
bands. Hindwing almost immaculate, slightly brownish
marginally, with blackish brown nygmata and similar
patterns of pterostigmatic markings. Veins brown, with
costal crossveins much darker. Rs seven-branched,
smoothly and straightly directed; three crossveins
between R1 and Rs; M two-branched; 1A two-branched.
Abdomen blackish brown with genital segments yel-
low. Ninth tergum (Fig. 13) broadly arcuate; anterior
margin deeply and arcuately concaved; posterior mar-
gin medially with a small arcuate incision. Tenth ter-
gum (Fig. 12) in lateral view subquadrate with round
posterior margin, moderately inflated in dorsal view.
Tenth sternum (Figs 12, 14–15) strongly sclerotized,
with short and rounded lateral arms; in ventral view
median plate ligulate, anteriorly incised V-shaped, pos-
terior half laterally with a pair of strongly elevated
ridges, and narrowed apex, which is slightly incised at
tip; in lateral view median plate moderately curved dor-
sad, posterior half ventrally with strongly elevated and
roundly margined ridge, and with tip narrowed.
Female. Unknown.
Type materials. Holotype Y, “C. Vietnam, Thua
Thien Hue, Bach Ma [16°12’N, 107°52’E], 1300 m,
2002.VI.8/11, M. Owada” (NSMT). Paratype: 1Y, same
data as holotype (NSMT).
Distribution. Vietnam (Thua Thien Hue Prov.).
Remarks. The new species is the single representa-
tive of the subfamily Chauliodinae in central Vietnam.
This species appears to be closely related to Neo-
chauliodes tamdaoensis sp. nov. in having the simi-
lar subquadrate male tenth tergum and the male tenth
sternum with elevated ridges. However, the new
species is much larger than N. tamdaoensis in body
size and the marking pattern of the forewing is also dif-
ferent from that of N. tamdaoensis. Considering the
genitalic structures, the new species can be distin-
guished from N. tamdaoensis by the male ninth ter-
gum with the medially incised posterior margin and the
male tenth sternum with the ventral ridges strongly
elevated, while in N. tamdaoensis the male ninth ter-
gum possesses the posterior margin without median
incision and the ventral ridges of the male tenth ster-
num is feebly elevated.
Neochauliodes bicuspidatus Liu et Yang
(Figs 2, 26)
Neochauliodes bicuspidatus Liu et Yang, 2006: 188. Type locality:
Yunnan (Longling, Wanding).
112 X. LIU, F. HAYASHI and D. YANG
Specimens examined. Holotype Y, “Yunnan,
Longling [a locality in Yunnan Prov., China,
24°35’N, 98°41’E], 1600 m, 1955.V.20, Le Wu” (IZCAS).
7Y2X, VIETNAM: no detailed data, 1995.V.10/17
(NSMT); 1Y, VIETNAM: Lai Chau Prov., Hoang
Lien Son Mts., N rdg., Mt. Phan [Fan] Si Pang
[22°20’N, 103°10’E], 1950–2050 m, 1995.V.17, A. Shino-
hara (NSMT); 1Y, VIETNAM: Lao Cai Prov., Fan
Si Pang Mts., Sa Pa [22°20’N, 103°49’E], 1995.III.25/30,
W. Mey (MFN); 1Y, “Yunnan [a locality in Yunnan
Prov., China, without precise data], Coll. Lacroix”
(MNHN).
Distribution. China (Yunnan Prov.); Vietnam (Lai
Chau Prov., Lao Cai Prov.).
Remarks. This species belongs to the Neochau-
liodes sinensis species group and is recorded for the
first time in Vietnam. For detailed description, see Liu
and Yang (2006).
Neochauliodes bowringi (McLachlan)
(Figs 3, 25)
Chauliodes bowringi McLachlan, 1867: 260. Type locality: Hongkong.
Specimens examined. Holotype Y, “Hongkong”
(NHM). 1X, VIETNAM: Ninh Binh Prov., Gia Vien, Cuc
Phuong [20°19’N, 105°37’E], 160 m, 1995.V.27, S. Ueno
(NSMT); 1X, VIETNAM: Vinh Phu Prov., Tam Dao
[21°45’N, 105°64’E], 1995.IV.17, Sinyajev (MFN); 1X,
“Tonkin Centr, Env. De Yen-bai [a locality in Yen Bai
Prov., Vietnam, without precise data], 1901, A. Weiss”
(MNHN); 1Y, “Kouy-Tchéou [a locality in Guizhou
Prov., China, without precise data], 1912, Père
Esquirol” (MNHN); 1Y2X, “China, Canton [a locality
in Guangdong Prov., China, without precise data],
Lehmann S.” (1Y in RMNH, 2X, in MFN); 1Y8X,
“China, Canton [a locality in Guangdong Prov.,
China, without precise data], Mell S.V.” (MFN); 1X,
“Lilong (China) [a locality in China without precise
data], Mus. Tuling” (MFN); 4Y1X, CHINA: Hong-
kong, Lantau I., San Shek Wan [22°12’N, 113°55’E],
1988.V, C. O’Connell (BPBM); 1Y, CHINA: Hongkong,
Lantau I., Nyong Ping, 440 m, 1987.IV.9, D. Batchelder
(BPBM).
Distribution. China (Fujian Prov., Guangdong
Prov., Guangxi Prov., Guizhou Prov., Hainan Prov.,
Hongkong, Hunan Prov., Jiangxi Prov., Shaanxi Prov.);
Vietnam (Ninh Binh Prov., Vinh Phu Prov., Yen Bai
Prov.).
Remarks. This species is a member of the Neo-
chauliodes bowringi species group and widely
distributed in central and southeastern China as well
as in northern Vietnam. For detailed description, see
Liu and Yang (2005a).
Neochauliodes confusus sp. nov.
(Figs 4, 16–19, 26)
Type locality. Vietnam (Tam Dao).
Etymology. The specific epithet “confusus” refers
to the undistinguished appearance of the new species,
which may cause confusion with its related species.
Description. Male. Body length 18–24 mm; fore-
wing length 29–31 mm, hindwing length 24–27 mm.
Head orange throughout. Compound eyes grayish
brown, ocelli yellow, medially margined black. Antenna
blackish brown with scape and pedicle brown. Mouth-
parts yellowish brown; mandibles with tips reddish
brown; maxillary and labial palpi with distal two seg-
ments blackish brown.
Prothorax orange; meso- and metathorax pale
brown, dorsally blackish brown laterally. Legs pale
brown with short, dense, brownish setae; tibiae and
tarsi blackish brown; tarsal claws reddish brown.
Wings hyaline with many brownish marks; pterostig-
matical area short, pale yellow. Forewing proximally
with a small spots on costal area, and with a short
stripe proximally to pterostigmatical area; proximal
markings small, rounded, densely dotted; median
transverse band narrow, extending to Cu2; distal
markings small, mostly dotted along longitudinal
veins. Hindwing similarly marked, with proximal half
entirely hyaline; median transverse band narrow,
extending to Cu1; distal markings usually fused with
each other along longitudinal veins. Veins yellow,
slightly darker on apex; costal crossveins blackish
brown. Rs six or seven-branched, posterior second and
third veins strongly curved posteriorly; three cross-
veins between R1 and Rs; M two-branched; 1A two-
branched.
Abdomen blackish brown. Tenth tergum (Fig. 16) in
lateral view subquadrate, about 1.5 times as long as
ninth tergum, with posterior margin nearly truncate; in
dorsal view apex distinctly inflated. Tenth sternum
(Figs 16, 18) strongly sclerotized, with short and subtri-
angular lateral arms; in ventral view median plate
attenuate with tip moderately narrowed, anterior mar-
gin arcuately incised, posterior margin slightly incised;
in lateral view median plate robust and curved dorsad
on posterior half, with tip obtuse and slightly nar-
rowed.
Female. Body length 22–29 mm; forewing length
31–38 mm, hindwing length 28–34 mm.
Eighth sternum (Fig. 19) subtrapezoidal in lateral
view, posterior margin strongly produced. Tenth
tergum short, subtriangular in lateral view, with
posterodorsal corner narrowed and produced. Ninth
gonocoxite broadly foliate.
Type materials. Holotype Y, “N. Vietnam, Vinh
Phu Prov., Tam Dao, 1230 m, 1995.IV.30/.V.1, M. Owa-
da” (NSMT). Paratypes: 1X, same data as holotype
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA
(NSMT); 1Y, VIETNAM: Vinh Phu Prov., Tam Dao,
1995.IV.21 (NSMT); 2X, VIETNAM: Vinh Phu Prov.,
Tam Dao, 1995.IV.22 (NSMT); 1Y, VIETNAM: Cao Bang
Prov., Mt. Pia Oac [22°37’N, 105°52’E], 1998.V.22/28, K.
Matsumoto (CAU); 1Y, CHINA: Guangxi Prov.,
Shangsi, Pinglongshan [22°09’N, 107°59’E], 350–500 m,
2002.IV.6, A. Shi (HBU).
Distribution . China (Guangxi Prov.); Vietnam
(Cao Bang Prov., Vinh Phu Prov.).
Remarks. The new species is a member of the
N. sinensis species group and only occurs in southern
Guangxi, China and northeastern Vietnam. This
species may be easily confused with Neochauliodes
punctatolosus Liu et Yang and Neochauliodes
formosanus (Okamoto) by the orange head and
pronotum, and the wings dotted by a great number of
small brownish spots. However, based on the male gen-
italic structures, it differs from the latter two species
by the male tenth sternum with posterior half of the
median plate not inflated in lateral view, and it seems
to be closely related to Neochauliodes meridionalis
van der Weele by the attenuate median plate of the
male tenth sternum. The broader male tenth tergum
and the robust median plate of the male tenth sternum
of the new species distinguis it from N. meridionalis.
Neochauliodes koreanus van der Weele
(Figs 5, 26)
Neochauliodes koreanus van der Weele, 1909: 261. Type locality:
Hongkong and Korea? (Seoul).
Specimens examined. Lectotype X, “Korea,
Seoul, 1900-251., August, Hon. E. Scarlett [probably a
wrongly attached label to the specimen]” (NHM).
7Y2X, VIETNAM: Vinh Phu Prov., Tam Dao,
1995.V.19/21 (NSMT); 3X, VIETNAM, Vinh Phu Prov.,
Tam Dao, 930 m, 1995.V.20, M. Sato (EU); 1X, VIET-
NAM: Vinh Phu Prov., Tamdao, 1993.V.18, K. Matsumo-
to (CAU); 1X, VIETNAM: Vinh Phu Prov., Tamdao,
1993.V, H. Karube (CAU); 1X, “China, Hongkong,
1908.V.24, F. Terry” (BPBM).
Distribution. China (Guangdong Prov., Guangxi
Prov., Hong Kong); Vietnam (Vinh Phu Prov.).
Remarks. This species belongs to the N. sinensis
species group and is distributed in southern China and
northern Vietnam. Its distribution in Korea is probably
erroneus. For detailed description, see Liu et al. (2007).
Neochauliodes meridionalis van der Weele
(Figs 6, 26)
Neochauliodes sinensis meridionalis van der Weele, 1909: 260.
Type locality: Vietnam.
113
Specimens examined. Lectotype X, “Haut-Tonkin
et Bas-Yunnan, Entre Man-Hao, Muong-Mum (Prés-
Lao-Kay) et Ban-Mam-Coun [a locality in Lao Cai
Prov. or Lai Chau Prov., Vietnam], 1905, Lieut Le-
sourt” (RMNH). 1X, “Tonkin, Rég. De Hoa-binh
[a locality in Hoa Binh Prov., Vietnam, without precise
data], 1928, A. De Cooman” (MNHN); 1X, “Tonkin, Ré-
gion de Hoa-binh” (MNHN); 1Y, “Tonkin, Région de
Hoa-binh, 1936, A. De Cooman” (MNHN); 1Y, CHINA:
Guangdong Prov., Chebaling [24°58’N, 114°02’E], 2001
(CAU); 6Y2X, CHINA: Guangdong Prov., Xinyi, Dawul-
ing [22°22’N, 110°55’E], 2002.V.31/ VI. 2 (SYSU); 1Y,
CHINA: Guangdong Prov., Zhaoqing, Dinghushan
[23°08’N, 112°29’E], 1964.VII.24, Z. Yang (SYSU).
Distribution. China (Guangdong Prov., Guangxi
Prov., Hainan Prov., Yunnan Prov.); Vietnam (Hoa Binh
Prov., Lai Chau Prov., Lao Cai Prov.).
Remarks. This species belongs to the N. sinensis
species group and is distributed in southern China and
northern Vietnam. Liu and Yang (2005a) elevated
N. sinensis meridionalis to a full specific status
based on the habitus photo given by van der Weele
(1910) and this species was revised by Liu et al. (2007).
However, at that time we did not examine the type
specimens. During our recent re-examination on the
historical megalopteran collections of several Euro-
pean museums, there are a great number of Neo-
chauliodes specimens found to be identified as
N. sinensis meridionalis by L. Navás and some
other early researchers, many of which are actually
not the true N. meriodionalis. Fortunately, we
found the two specimens noted as types (syntypes)
by van der Weele (1910) currently deposited in RMNH
and MNHN respectively. The specimen, for which the
habitus photo in the monograph of van der Weele
(1910) was taken, is a female specimen labeled “Haut-
Tonkin et Bas-Yunnan, Entre Man-Hao, Muong-Mum
(Prés-Lao-Kay) et Ban-Mam-Coun, 1905, Lieut Le-
sourt” but missing the type label. The other male spec-
imen in MFN is labeled “Mou-pin, 1901, R. Oberthür”
and bears a type label. After our re-examination,
the female syntype is identical to our concept of this
species, but the male specimen is actually Neo-
chauliodes sinensis (Walker). So, to prevent mak-
ing a new synonym, we designated the female spec-
imen in RMNH as the lectotype, and the male
specimen, which was misidentified as N. sinensis
meridionalis by van der Weele (1909), as the paralec-
totype.
Neochauliodes moriutii Asahina
(Figs 7, 25)
Neochauliodes moriutii Asahina, 1988. Gekkan-Mushi 207: 8. Type
locality: Thailand (Chiang Mai).
114 X. LIU, F. HAYASHI and D. YANG

Figures 1–3. Habitus photos of the Neochauliodes species from Indochina. (1) N. bachmanus sp. nov., male, holotype;
(2) N. bicuspidatus Liu et Yang, male from Vietnam; (3) N. bowringi (McLachlan), female from Vietnam. Scale bars = 5.0 mm
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA 115

Figures 4–6. Habitus photos of the Neochauliodes species from Indochina. (4) N. confusus sp. nov., male, holotype; (5) N. koreanus van
der Weele, male from Vietnam; (6) N. meridionalis van der Weele, female, lectotype. Scale bars = 5.0 mm
116 X. LIU, F. HAYASHI and D. YANG

Figures 7–9. (7) N. moriutii Asahina, male, holotype; (8) N. puntatolosus Liu et Yang, male from Laos; (9) N. tamdaoensis sp. nov.,
male, holotype. Scale bars = 5.0 mm.
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA 117

Specimens examined. Holotype Y, “Thailand,
Chiang Mai, Doi Pui [19°39’N, 99°45’E], 1985.V.18”
(NSMT). Allotype X, “N. Thailand, Doi Pui, 1985.V.19”
(NSMT). Paratypes: 1Y, THAILAND: Chiang Mai Prov.,
Doi Chang Khian [18°49’N, 98°57’E], 1981.VII.21, H.
Kuroko, S. Moriuti, Y. Arita & Y. Yoshiyasu (NSMT);
2Y, THAILAND: Chiang Mai Prov., Doi Inthanon (Mae
Klang) [18°39’N, 98°25’E], 1987.IX.8/12, S. Moriuti, Y.
Arita & Y. Yoshiyasu (NSMT). 7Y, THAILAND: Chiang
Mai Prov., Doi Inthanon (Mae Klang), 1987.IX.8/12, S.
Moriuti, Y. Arita & Y. Yoshiyasu (NSMT); 6Y, LAOS:
Houaphan Prov., Xamneua, Mt. Phu Pan Saluei
[20°25’N, 104°02’E], 2003.V (EUM); 1Y, LAOS: Hou-
aphan Prov., Xam Neua around Ban Saleui, 2004.IV.9/
10, N. Ohbayashi (EUM); 2Y, LAOS: Vientiane Prov.,
Ban Van Heue, Phou-kow-kwei [18°48’N, 102°35’E], 800
m, 1965.IV.14/16, J.L. Gressitt (BPBM); 1X, “Carin
Chebà [a locality in Kayin State, Myan-mar, 19°13’N,
96°35’E], 900–1100 m, VI.1888, L. Fea” (RMNH); 1X,
THAILAND: Changwat Nan [Nan Prov.], 30 km E of
Pua [19°13’N, 100°54’E], 1700 m, 1998.II.27, M. Hreblay
& C. Szabóky (HNHM).
Distribution. China (Yunnan Prov.); Laos (Houa-
phan Prov., Vientiane Prov.); Myanmar (Kayin Stat.);
Thailand (Chiang Mai Prov., Nan Prov.).
Remarks. This species is a member of the N. bow-
ringi species group. Besides its type locality in
Thailand, Liu and Yang (2006) recorded this species
in China (southern Yunnan Prov.) for the first time
and made a redescription. Herein, we add Laos
and Myanmar as the new country records of this
species.

10

11

Figures 10–11. (10) N. tonkinensis (van der Weele), female, holotype; (11) N. umbratus Kimmins, male from Vietnam.
Scale bars = 5.0 mm.
118 X. LIU, F. HAYASHI and D. YANG

Neochauliodes punctatolosus Liu et Yang
(Figs 8, 26)
Neochauliodes punctatolosus Liu et Yang, 2006: 193. Type locality:
Yunnan (Jingdong).
Specimens examined. Holotype Y, “Yunnan, Jing-
dong [a locality in Yunnan Prov., China, 24°26’N,
100°50’E], 1170 m, 1956.VII.4” (CAU). 1X, VIETNAM:
Son La Prov., Truong Yen [20°50’N, 104°38’E], 1995.V.1,
A. Saito (NSMT); 2Y, THAILAND: Chiang Mai Prov.,
Chiang Dao [19°20’N, 98°57’E], 2004.V.12, I. Sivec & H.
Bogdan (CAU); 5Y, THAILAND: Chiang Mai Prov.,
Fang [19°55’N, 99°12’E], 450 m, 1983.V.14/17, H.
Kuroko, S. Moriuti, Y. Arita & Y. Yoshiyasu (NSMT);
1Y, THAILAND: Mae Sa Water-fall [a locality in Chi-
ang Mai Prov., 18°54’N, 98°54’E], 1965.VI.16, S. Asahina
(NSMT); 1X, THAILAND: Chiang Mai Prov., Ta Ton
[19°30’N, 99°04’E], 1981.VII.17, H. Kuroko, S. Moriuti, Y.
Arita & Y. Yoshiyasu (NSMT); 1Y, THAILAND: Chan-
thaburi Prov., Khao Soi Dan [12°54’N, 102°12’E], 400 m,
1983.VI.6, H. Kuroko, S. Moriuti, Y. Arita & Y. Yoshiya-
su (NSMT); 1X, “Laos, Rég de Xiang-khouang Tran-
ninh [a locality in Xieng Khoang Prov., Laos, without
detailed data], 1.200 A 1200 m, 1920, R. Miéville”
(MNHN); 1X, “Laos, Xieng-kouang [a locality in Xieng
Khoang Prov., Laos, without detailed data], 1922,
Colonico” (MNHN); 1Y, LAOS: Houaphan Prov., Ban
Saleui N.E., Mt. Pan (CAU); 1Y, LAOS: Louang Namtha
Prov., Muong Sing [21°11’N, 101°08’E], 650 m,
1960.VI.8, L.W. Quate (BPBM); 1X, LAOS: Vientiane

12 13

14

15

Figures 12–15. Neochauliodes bachmanus sp. nov. (12) Male genitalia, lateral view; (13) male genitalia, dorsal view; (14) male tenth
sternum, ventral view; (15) male tenth sternum, caudal view. Scale bars = 1.0 mm.
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA
Prov., Ban Van Heue, 1965.IV.13, Gressitt (BPBM);
4Y2X, LAOS: Houaphan Prov., Xamneua, Mt. Phu Pan
Saluei, 2003.V (EUM).
Distribution. China (Yunnan Prov.); Laos (Houa-
phan Prov., Louang Namtha Prov., Vientiane Prov.,
Xieng Khoang Prov.); Thailand (Chanthaburi Prov.,
Chiang Mai Prov.); Vietnam (Son La Prov.).
Remarks. This species belongs to the N. sinensis
species group and is distributed in southern Yun-
nan, northern Laos, Thailand, and northwestern
Vietnam. For detailed description, see Liu and Yang
(2006).
Neochauliodes tamdaoensis sp. nov.
(Figs 9, 20–25)
Type locality. Vietnam (Tam Dao).
Etymology. The specific epithet “tamdaoensis”
refers to the type locality of the new species.
Description. Male. Body length 18–19 mm; fore-
wing length 29–32 mm, hindwing length 27–28 mm.
Head yellowish brown, frons blackish brown and
somewhat extended toward vertex. Compound eyes
dark brown, ocelli yellow, medially margined black.
Antenna blackish brown. Mouthparts dark brown.
16
18
Figures 16–19. Neochauliodes confusus sp. nov. (16) Male genitalia, lateral view; (17) male tenth tergum, dorsal view; (18) male
tenth sternum, ventral view; (19) female genitalia, lateral view. Scale bars = 1.0 mm.
119
Thorax pale brown. Legs brown with short, dense,
brownish setae; tarsal claws reddish brown. Wings
hyaline with many brownish markings; pterostigmati-
cal area short, pale yellow. Forewing with several
small spots on proximal half of costal area, and respec-
tively with a long stripe proximally and a short stripe
distally to pterostigmatical area; basal marks small,
dispersed dotted; median markings generally formed
into three narrow transverse bands, distal one of which
somewhat separated; distal margin mostly pale brown.
Hindwing similarly marked except for proximal half
entirely hyaline, with only one short median transverse
band extending to M. Veins brown. Rs seven or eighth-
branched, almost straightly directed; three cross-
veins between R1 and Rs; M two-branched; 1A two-
branched.
Abdomen dark brown. Tenth tergum (Fig. 20) in
lateral view subquadrate with rounded posterior
margin; in dorsal view short, strongly inflated. Tenth
sternum (Figs 22–23) strongly sclerotized, with short
and rounded lateral arms; in ventral view median
plate attenuate, anterior margin widely incised,
V-shaped, posterior half laterally with a pair of
feebly elevated ridges, posterior 1/3 distinctly nar-
rowed with rounded tip; in lateral view median
plate feebly curved dorsad, posterior half ventrally
17
19
120 X. LIU, F. HAYASHI and D. YANG

with feebly elevated ridge, and with tip strongly nar-
rowed.
Female. Body length 26 mm; forewing length 39 mm,
hindwing length 34 mm. Wings with markings much
darker and broader than those in male.
Eighth sternum (Fig. 24) subtrapezoidal in lateral
view, posterior margin strongly produced. Tenth
tergum shortly claviform, moderately produced dor-
sad and slightly narrowed toward tip; cercus large,
strongly inflated, slightly wider than apex of tenth
tergum. Ninth gonocoxite elongated, subtriangular,
slightly produced dorsad, proximal half mostly sclero-
tized.
Type materials. Holotype Y, “Vietnam, Vinh Phu
Prov., Tam Dao, 1995.V.20, M. Owada” (NSMT).
Paratypes: 1X, same data as holotype (NSMT); 1Y,
VIETNAM: Vinh Phu Prov., Tam Dao, 960–1230 m,
1995.V. 19/21, A. Shinohara (NSMT).
Distribution. Vietnam (Vinh Phu Prov.).
Remarks. See remarks of N. bachmanus.
Neochauliodes tonkinensis (van der Weele)
(Figs 10, 25)
Chauliodes tonkinensis van der Weele, 1907: 260. Type locality: Viet-
nam (Chiem-Hoa).
Neochauliodes orientalis Yang et Yang, 1991: 74. Type locality: Yun-
nan (Binchuan, Huaning, Kunming, Changning, Gejiu). syn. nov.
Neochauliodes yunnanensis Navás, 1930: 1. Type locality: Yunnan
(San-heu-kai). syn. nov.
Specimens examined. Holotype X, “Central-
Tonkin, Chiem-Hoa [a locality in Tuyen Quang Prov.,
Vietnam, 22°15’N, 105°15’E], Aug. Sept., H. Fruhstor-
fer” (RMNH). 1Y [Holotype of Neochauliodes orien-
talis Yang & Yang], “Yunnan, Binchuan [a locality in

20 22

24
21

23

Figures 20–24. Neochauliodes tamdaoensis sp. nov. (20) Male genitalia, lateral view; (21) male tenth tergum, dorsal view; (22) male
tenth sternum, ventral view; (23) male tenth sternum, caudal view; (24) female genitalia, lateral view. Scale bars = 1.0 mm.
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA 121

Yunnan Prov., China, 25°49’N, 100°32’E], 1800 m, 1980.
VI.9” (CAU); 1Y [Holotype of Neochauliodes yunna-
nensis Navás], “Yunnan, San-heu-kai [a locality in
Yunnan Prov., China, without precise data]” (MZBS);
2X, “Yunnan, Coll. Lacroix” (MNHN); 5Y, VIETNAM:
Cao Bang Prov., Mt. Pia Oac, 1985.V.14 (HC); 1X, VIET-
NAM: Vinh Phu Prov., Tam Dao, 950 m, 1999.V. 14/17,
M. Owada (NSMT); 34Y4X, VIETNAM: Vinh Phu Prov.,
Tam Dao, 1995.V.13/23 (NSMT); 5Y1X, VIETNAM: Lao
Cai Prov., Sa Pa, Deo O Quy Ho, 1750 m, 1999.VI. 1/3, M.
Owada (NSMT); 10Y7X, VIETNAM: Lao Cai Prov., Fan
Si Pang Mts., Sa Pa, 1995.III.25/30, W. Mey (MFN);
4Y2X, VIETNAM: Lao Cai Prov., Fan Si Pang Mts.,
Sa Pa, 1600 m, 1995.IV.22/29 (MFN); 1Y, “Hte. Bir-
manie, Etatde Momeit [Möng Mit, in Shan State, Myan-
mar, 23°06’N, 96°40’E], 600 m, 1890, Doherty” (MNHN);
1Y, LAOS: Houaphan Prov., Ban Saluei, 1400–1500 m,
2003. VI.20/24, N. Ohbayashi (EUM); 1Y, LAOS: Houa-
phan Prov., Xamneua, Mt. Phu Pan Saluei, 2003.V
(EUM).
Distribution. China (Yunnan Prov.); Laos (Houa-
phan Prov.); Myanmar (Shan Stat.); Vietnam (Cao
Bang Prov., Lao Cai Prov., Tuyen Quang Prov.; Vinh
Phu Prov.).
Remarks. This species is a member of the N. bow-
ringi species group. It resembles N. moriutii by the
similar pattern of the wings. However, compared with
N. moriutii, the markings on the distal half of the
wings are usually much darker and denser in N. tonk-
inensis. The male tenth sternum of N. tonkinensis is
gradually tapering posteriorly, while it is nearly subob-
long with truncate tip in N. moriutii.
After checking the holotype of N. tonkinensis,
we found that Neochauliodes orientalis Yang
et Yang, which was redescribed by Liu and Yang
(2006), is conspecific with the former species and
should be treated as a junior synonym. Additionally,
Neochauliodes yunnanensis Navás is also treated as
a junior synonym of N. tonkinensis by the kind help of
Dr. V. Monserrat for checking holotype.

Figure 25. Distribution map of Neochauliodes spp. from Indochina. (
) N. bachmanus sp. nov.; () N. bowringi (McLachlan);
() N. moriutii Asahina; (
) N. tamdaoensis sp. nov.; () N. tonkinensis (van der Weele); (+) N. umbratus Kimmins.
122 X. LIU, F. HAYASHI and D. YANG

Neochauliodes umbratus Kimmins na, Canton [Guangdong Prov.], Ting-wu-san [Dingh-

(Figs 11, 25)
Neochauliodes umbratus Kimmins, 1954: 436. Type locality: Indo-
China.
Neochauliodes furcatus Yang et Yang, 1986: 91. Type locality:
Guangxi (Longzhou).
Specimens examined. Holotype Y, “Indo-China
[a locality probably in northern Vietnam], 1933-222,
A. Vuillet” (NHM). 1Y1X, “Tonkin, env. d. Hanoi
[21°01’N, 105°51’E], E. Le Moult” (NSMT); 1Y, VIET-
NAM: Ha Tay Prov., Ao Vua [21°05’N, 105°22’E], 1995.
IV.27 (NSMT); 4Y6X, VIETNAM: Vinh Phu Prov., Tam
Dao, 1995.IV.20/23 (NSMT); 3Y, VIETNAM: Vinh Phu
Prov., Tam Dao, 1230 m, 1995.IV.30/V.1, M. Owada
(NSMT); 1Y1X, VIETNAM: Vinh Phu Province, Tam
Dao, 1200 m, 1999.V.24/25, H. Karube (HC); 2X, VIET-
NAM: Vinh Phu Province, Tam Dao, 1998.V, K. Ikeda
(HC); 1Y, VIETNAM: Vinh Phu Prov., Tam Dao,
1995.IV.17, Sinyajev (MFN); 1X, VIETNAM: Vinh Phu
Prov., Tam Dao, 1995.IV.22, M. Sato (EUM); 1X, “Chi-
ushan], Mell S.V.” (MFN).
Distribution. China (Guangdong Prov., Guangxi
Prov.); Vietnam (Hanoi, Ha Tay Prov., Vinh Phu
Prov.).
Remarks. This species is rather distinct in Neo-
chauliodes by the male tenth sternum with addition-
al processes. Liu and Yang (2005a) redescribed this
species based on material from China. The record of
this species in Indochina had been obscure due to the
single inaccurate collecting label of the holotype. Here-
in, based on the finding of more material from Vietnam,
the range of this species is generally around Hanoi, Ha
Tay Prov., and Vinh Phu Prov. in Vietnam and south-
eastern China.
DISCUSSION
The list of Neochauliodes from Indochina now
contains 11 species. Three species, N. simplex,

Figure 26. Distribution map of Neochauliodes spp. from Indochina. () N. bicuspidatus Liu et Yang; (
) N. confusus sp. nov.;
(+) N. koreanus van der Weele; (
) N. meridionalis van der Weele; () N. punctatolosus Liu et Yang.
 THE GENUS NEOCHAULIODES VAN DER WEELE FROM INDOCHINA
N. sinensis (formerly N. sinensis sinensis), and
N. occidentalis (formerly N. sinensis occidentalis),
presented in the species list from Indochina (Lestage
1927), are herein excluded from this area. The speci-
men identified as N. simplex from Kouy-Tchéou
Indochina (MNHN), which was considered to be this
species by Lestage (1927), actually belongs to N. bow-
ringi based on our examination. Similarly, the speci-
men from Yunnan (MNHN), the basis for which N. si-
nensis was recorded from Indochina by Lestage
(1927), is N. bicuspidatus. Although we did not exam-
ine the specimens listed by Lestage (1927) as N. occi-
dentalis from Laos and Vietnam, the occurrence of
this species in Indochina is quite doubtful. The true
N. occidentalis is currently known only in Sichuan
Prov. and Gansu Prov. of west China (Liu et al. 2007),
and is not found in south China and adjacent areas
in Indochina based on our recently intensive inves-
tigation.
The Neochauliodes fauna of Indochina is distinct
from those of the Indian subcontinent, Malaysia, and
Indonesia, without any common species among these
areas. However, nine species of Neochauliodes are
distributed both in Indochina and mainland China,
showing a close faunistic relationship between these
two regions. Within the 11 Neochauliodes species in
Indochina, 10 species are distributed in Vietnam, two of
which are endemic to this country, while only three
species occur in Myanmar, Thailand, and Laos, and
none are recorded in Cambodia. Thus, northern Viet-
nam is considered to the center of diversity of
Neochauliodes in Indochina, with nine species res-
tricted to several mountain ranges of this area, includ-
ing Hoa Lien Son, Moc Chau Plateau, Ba Vi, Tam Dao
and Mt. Pia Oac. However, the current diversity of
Neochauliodes in Indochina is less than our expecta-
tion. The collections of Megaloptera are still lacking in
vast area of Indochina, especially in Myanmar, Laos,
and Cambodia. More new discoveries from Indochina
are expected in the future when more intensive collec-
tion are made.
ACKNOWLEDGEMENTS
We are very grateful to the following colleagues for
their kind help granting us access to the megalopteran
specimens in their collections: Prof. C. van Achterberg
(Leiden), Dr. R. de Vries (Leiden), Prof. M. Ohl (Berlin),
Prof. J. Legrand (Paris), Dr. G. Sziráki (Budapest),
Dr. D. Goodger (London), Dr. V. Monserrat (Madrid),
Dr. N.L. Evenhuis (Honolulu), Dr. M. Owada (Tokyo),
Prof. N. Ohbayashi (Matsuyama), Mr. S. Matsuno
(Matsuyama), Prof. H. Pang (Guangzhou), Prof. G. D.
Ren (Baoding), Mr. J. Yao (Beijing), and Ms. H. Liu
123
(Beijing). This research was supported by a grant-in-
aid for fellows of the Japan Society for the Promotion of
Science (JSPS) relating to the JSPS Postdoctoral Fel-
lowship for Foreign Researchers (No. 20-08417) and by
the Science Foundation of Doctor Subjects, State Edu-
cation Ministry of China (No. 20090008120019).
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