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ACTA ENTOMOLOGICA
MUSEI NATIONALIS PRAGAE
www.aemnp.euISSN 1804-6487 (online) – 0374-1036 (print)
RESEARCH PAPER
Review of the genus Cybister in China, with description
of a new species from Guangdong (Coleoptera: Dytiscidae)
Zhuo-Yin JIANG1), Shuang ZHAO2), Zu-Qi MAI3), Feng-Long JIA4,6) & Lars HENDRICH5,6)
1) Institute of Entomology, Life Science School, Sun Yat-sen University, Guangzhou 510275, China; e-mail: zhuoyin_jiang@126.com;
ORCID ID https://orcid.org/0000-0001-6825-4283
2) Guangzhou Customs Technology Center, Tianhe District, Guangzhou, China; e-mail: topzs@163.com;
ORCID ID https://orcid.org/0000-0002-0963-6879
3) School of Agriculture, Sun Yat-sen University, Guangzhou, 511436, China; e-mail: maizq5@mail2. sysu.edu.cn;
ORCID ID https://orcid.org/0000-0003-3124-2021
4) Institute of Entomology, Life Science School, Sun Yat-sen University, Guangzhou 510275, China;
e-mail: lssjfl @mail.sysu.edu.cn, fenglongjia@aliyun.com; ORCID ID http://orcid.org/0000-0003-2391-5038
5) Department of Entomology, SNSB-Zoologische Staatssammlung München, München, Germany;
e-mail: hendrich@snsb.de; ORCID ID https://orcid.org/0000-0002-3773-6586
6) corresponding author
Abstract. The genus Cybister Curtis, 1827 is reviewed with respect to the fauna of China. A
new species, Cybister (Melanectes) danxiaensis sp. nov. is described and illustrated. Altogether,
16 Cybister species are recorded from China, although the status of two of them remains in
question. All species are diagnosed, with their habitus and male genitalia illustrated. Additi-
onal faunistic data from China are provided for the following species: C. guerini Aubé, 1838
from Hainan; C. lewisianus Sharp, 1873 from Guangdong and Hainan; C. limbatus (Fabri-
cius, 1775) from Guangdong, Guangxi and Hainan; C. rugosus (W. S. Macleay, 1825) from
Chongqing, Guangdong, Hubei, Hunan, Jiangxi, Yunnan, Zhejiang; C. tripunctatus lateralis
(Fabricius, 1798) from Chongqing, Guangxi, Jiangxi, Shanghai; C. brevis Aubé, 1838 from
Guizhou, Hunan, Zhejiang; and C. sugillatus Erichson, 1834 from Guangxi and Yunnan.
An updated identifi cation key and distribution maps for all species are provided. Cybister
lateralimarginalis torquatus (Fischer von Waldheim, 1829) is recorded for the fi rst time from
Tajikistan and Uzbekistan.
Key words. Coleoptera, Dytiscidae, Cybistrinae, Cybister, taxonomy, new species, new re-
cords, China, Palaearctic Region
Zoobank: http://zoobank.org/urn:lsid:zoobank.org:pub:937DF12A-BC2F-4406-802F-63838312B0C1
© 2023 The Authors. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Licence.
Accepted:
21st February 2023
Published online:
12th March 2023
2023
63(1): 75–102
doi: 10.37520/aemnp.2023.003
Introduction
Cybister Curtis, 1827 is the only genus of Cybistrinae
that occurs in China. It comprises a total of 106 known
species/subspecies worldwide (N & H 2022a).
These beetles are large to extremely large (13–43 mm),
dark green to black, and in some species with yellow
margins of the pronotum and elytra. Members of the
genus typically inhabit stagnant waters, generally with
dense submerged aquatic vegetation. Although species of
Cybister occur in all zoogeographical regions, the genus
is most abundant in the Afrotropical and Oriental Realms
(M & B 2016).
S (1882) divided Cybister species into six groups
based on morphological characters, and B (1945)
divided the genus into seven subgenera. However, V-
(1969) discovered some problems with the classi-
fi cation of the groups and subgenera when he studied the
male genitalia of Indian Cybister. M et al. (2007)
re-evaluated the classifi cation of the genus and proposed
four subgenera based on the results of both morphological
and DNA analysis.
In China, there were few studies on Cybister until F
(1932, 1933a) recorded ten species in his catalogue of
Chinese Dytiscidae. Based on the material he studied and
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
76
the data from Z (1920), F (1933b) pro-
vided a detailed faunistic report with a key in which eight
Cybister species were included. W (1937) recorded ten
species in China. N (1995) reported sixteen species
in his annotated check-list of China based on a literature
review. J et al. (2010) provided faunistic data on Cybis-
ter in China in which two species names were treated as
synonyms and one species was recorded as new for China.
Up to now, a total of fi fteen species have been recorded
from China (J et al. 2010, N & H 2022b),
but a comprehensive taxonomic revision of Chinese or
continental Asian species is still missing. Therefore, the
identifi cation of some species (e.g. Cybister guerini Aubé,
1838, C. limbatus (Fabricius, 1775)) treated in the present
publication can be tentative only. Additional information
about Cybister species occurring in China can be found
also in the Indian revision by V (1969), or in the
faunistic review of Cybister from Laos, including habitus
and habitat photos by H B (2013).
In the present work, we describe another new species in
the subgenus Melanectes Brinck, 1945 from northern Gu-
angdong, increasing the total number of Chinese Cybister
to 16. In addition, we summarize all previously published
records and provide numerous new faunistic data for some
species. For the fi rst time, the habitus and male genitalia
are illustrated for all species occurring in China, and an
identifi cation key is provided.
Material and methods
The studied specimens were examined and measured
under a Nikon SMZ800N binocular microscope and a
Leica M205C stereomicroscope. Male specimens of each
species were dissected. First relaxed in hot water for 30
minutes and then the genital capsule extracted with strong,
pointed forceps with curved tips. The median lobe of the
aedeagus and the parameres were glued to a small card for
photographing, and mounted together with the specimen
after being photographed. Photographs of habitus, detailed
characteristics and male genitalia were taken with a Nikon
DS-Ri2 camera mounted on a Nikon SMZ25 microscope
and a Canon EOS 550D camera fi tted with either a 65 mm
or MPE65 macro lens, attached to a Stackshot Macrorail
controlled with Zerene Stacker software; Illumination was
with two Canon Speedlite 430EX III-RT fl ashlights and
translucent paper diff usors. Images were assembled using
Helicon Focus and NIS-Elements software and were edited
with Adobe Photoshop CS6.
The following abbreviations are used in the descriptions:
TL total length, measurement of length from clypeal margin to
apex of elytra;
TL-h total length minus head length, measurement of length from
anterior margin of pronotum to apex of elytra;
MW maximum width of body measured at right angle to TL.
The terminology used to describe morphological
structures follows M & B (2016). The ter-
minology to denote the orientation of the genitalia follows
M & N (2003). The species are listed alpha-
betically. Exact label data are cited for the type material
and given in quotation marks. Authors’ additional remarks
are provided in square brackets; [hw] – preceding data are
handwritten; [p] – preceding data are printed. Separate
label lines are indicated by a slash (/), separate labels by
a double slash (//).
The specimens included in this study are deposited in
the following collections:
JSCL Jaroslav Šťastný collection, Liberec, Czech Republic;
LHCM Lars Hendrich collection, München, Germany (property of
NHMW);
NHMB Naturhistorisches Museum, Basel, Switzerland;
NHRS Swedish Museum of Natural History, Stockholm, Sweden;
NMPC Národní muzeum, Prague, Czech Republic;
SYSU Biological Museum, Sun Yat-sen University, Guangzhou,
China;
ZMUC Zoological Museum, University of Copenhagen, Copenhagen,
Denmark;
ZSMG Zoologische Staatssammlung, München, Germany.
Taxonomy
Subgenus Cybister Curtis, 1827
Type species. Dytiscus lateralis Fabricius, 1798.
Diagnosis. With distinctive yellow margins laterally on
the pronotum and elytron. Males with a single claw per
metatarsus; females usually with single claws, except in
some North and Central American species.
Included (Chinese) species. Cybister bengalensis Aubé,
1838; C. chinensis Motschulsky, 1854; C. fumatus Sharp,
1882; C. guerini Aubé, 1838; C. laevis Falkenström, 1936;
C. lateralimarginalis torquatus (Fischer von Waldheim,
1829); C. lewisianus Sharp, 1873; C. limbatus (Fabricius,
1775); C. rugosus (W. S. Macleay, 1825); C. tripunctatus
lateralis (Fabricius, 1798); C. ventralis Sharp, 1882.
Cybister bengalensis Aubé, 1838
(Figs 1A–E)
Cybister bengalensis Aubé, 1838: 61 (orig. descr.).
Material examined. CHINA: J: 1 , Pingshiang [Pingxiang],
coll. A. Zimmermann (ZSMG). T: 2 , Formosa, coll. A. Zimmer-
mann (ZSMG); 1 , Ku Sia, coll. A. Zimmermann (ZSMG). W
: 1 , China, coll. A. Zimmermann (ZSMG).
Diagnosis. See under C. rugosus.
Comments to classifi cation. Almost all characters of C.
bengalensis described by R (1899) also appear
in C. rugosus. Based on the material we have studied from
South China , C. bengalensis could be conspecifi c with
C. rugosus as was suggested already by G
(1938) after he studied aedeagi of both species. The status
of C. bengalensis and C. rugosus should be confi rmed after
the type material of the two species will be studied in future.
Careful study of the specimens deposited in SYSU and
a few specimens deposited in the Institute of Zoology,
Chinese Academy of Sciences, Beijing, China, we found
the specimens of C. bengalensis, identifi ed by Tong-xu
Peng and checked by Zeng, from Guangzhou in SYSU are
mixed with specimens of C. guerini and C. rugosus. The
specimens from Beijing and Hebei should be C. chinensis;
the record from Northern Territory of China needs to be
confi rmed as well.
Distribution. Cybister bengalensis was originally de-
scribed from “East India and China”. It was subsequently
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 77
Fig. 1. Cybister bengalensis Aubé, 1838 (, Ku Sia, Taiwan). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus in
lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
recorded from North China (S 1882), Sichuan (S
1882, R 1899, Fö 1936), Jiangxi and
Guangdong (R 1899), and further from Beijing,
Fujian, Hainan (F 1932), Zhejiang (W 1937), Hebei
and Yunnan (Z 1989, N 1995, J et al. 2010).
Cybister chinensis Motschulsky, 1854
(Figs 2A–E; 18A–C; 19A; 20A)
Cybister chinensis Motschulsky, 1854: 44 (orig. descr.).
Cybister japonicus Sharp, 1873: 45 (orig. descr.), synonymy by Z
(1915): 284 and confi rmed by N & P (2007): 43; S
(1882): 748 (Manchuria, Shandong, Taiwan); R (1899): 348
(Manchuria, Taiwan); F (1932): 36 (Beijing, Fujian: Fuzhou and
Xiamen, Guangdong, Hainan, Manchuria, Shandong: Yantai, Taiwan);
F (1936): 244 (Sichuan); F (1936): 13 (Liaoning:
Shenyang, Shaanxi); G (1938): 76 (Manchuria,
Taiwan); B (1946): 151 (Heilongjiang, Liaoning); Z
(1972): 360 (Taiwan); Z (1981): 110 (Fujian); Z (1989): 90
(NE China, Fujian, Guangdong, Hainan, Hebei, Ningxia: Yinchuan,
Shandong, Taiwan); L (1992): 36 (Jilin, Liaoning); M & K
(1993): 145 (Taiwan).
Material examined. CHINA: G: 3 , Canton, 1.–21.
iii.2001, L. Hojný leg. (NMPC). N: 1 , Yinchuan, under light,
6.vi.1960 (SYSU); 1 , Yinchuan, under light, 15.vii.1962 (SYSU); 1
, Yinchuan, Gujiaqiao, 23.vi.2011 (SYSU); 2 , Yinchuan (SYSU).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
78
Fig. 2. Cybister chinensis Motschulsky, 1854 (, Heihe, Heilongjiang). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of
aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
T: 1 , Tsing tou, 17.viii.1933 (SYSU); 1 , Tientsin, 10.iv.1935
(SYSU); 1 , Tianjin, Balitai, 4.v.1955, Y. Y. Liu leg. (SYSU). W
: 15 spec., donated by Xing-min Wang, purchased from
market, without data (SYSU); 1 , China (LHCM).
Diagnosis. TL 35–40 mm, elongate, oblong-oval (Fig.
2A). Head dark green, clypeus yellow. Pronotum and
elytra dark green to black, with distinct yellow margins
laterally. Ventral surface yellow; prosternal process, mid-
dle of metaventrite and metacoxae dark green; metacoxae
and abdominal ventrites narrowly margined (Fig. 2B).
Dorsal surface dimorphic: smooth with sparse fi ne punc-
tures in male; in female with isolated short, irregular
striae on pronotum, and with dense longitudinal striae
of various lengths on elytra except at its apex. Prosternal
process lanceolate, with distinct lateral bead, surface fl at,
apex pointed; lateral parts of metaventrite (ʻmetasternal
wingsʼ) tongue-shaped, slender, not reaching lateral
margins; metacoxal lines distinct, well impressed, nearly
parallel medially, divergent anteriorly, along metacoxal
lines with a row of punctures. Forelegs yellow, midlegs
and hindlegs yellow to brown; protarsomeres 1–3 of male
broadly expanded into a palette, with four rows of adhe-
sive setae on their ventral side; metatarsal claws single in
both sexes. Median lobe of aedeagus ʻCʼ-shaped in lateral
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 79
view, moderately curved (Fig. 2C); ridged in middle in
ventral view, lateral margin strongly curved near apex,
apex obtuse-angled (Fig. 2D). Parameres moderately
broad, strap-like apically (Fig. 2E).
Habitat. This species inhabits ponds, paddy fi elds and
wetlands with aquatic plants (M & K 1993);
specimens from Shanxi were found in a natural lake (Figs
18A–C).
Distribution. China (Beijing, Fujian, Guangdong,
Hainan, Hebei, Heilongjiang, Jilin, Liaoning, Ningxia,
Sichuan, Shaanxi, Shandong, Shanxi, Taiwan, Tianjin)
(Fig. 20A), Japan, North Korea, Russia, South Korea
(R 1899; Z 1920; F 1932, 1933b;
G 1938; Z 1972; Z 1989; M &
K 1993; N 1995; N & Há 2022b).
Cybister fumatus Sharp, 1882
(Figs 3A–E; 20A)
Cybister fumatus Sharp, 1882: 731 (orig. descr.); F (1936): 13
(Shanghai).
Material examined. None from China but 9 specimens from LAOS: 4
spec., Kham Mouan prov., Ban Khoun Ngeun, 18°07′N 104°29′E, 200
m, 19.–31.v.2001, P. Pacholátko leg. (NHMB, LHCM); 1 spec., Vien-
tiane prov., Phou Khao Khouay, 18°20.369′N 102°48.523′E, nr. strongly
disturbed primary rainforest on light, 25.–30.v.2008, A. Solodovnikov
& J. Pedersen leg. (ZMUC); 1 spec., Sekong Prov., Tad Faek waterfalls,
15°14.7′N 106°42.1′E, 118 m, light trap, 8.v.2010, J. Hájek leg. (NMPC);
Fig. 3. Cybister fumatus Sharp, 1882 (, Laos). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus in lateral view;
D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
80
1 spec., Bolikhamsay prov., Nam Kading NPA research center near
Ban Phone Kham, 18°20′N 104°08′E, 250 m, 23.–29.v.2011, NHMB
Basel Laos 2011 Expedition M. Brancucci, M. Geiser, D. Hauck, Z.
Kraus, A. Phantala & E. Vongphachan (NHMB); 1 spec., Bolikhamsay
prov., Pakkading Ban Phone Kham env. 18°19′N 104°08′E 200–300 m,
23.–29.v.2011, NHMB Basel Laos 2011 Expedition M. Brancucci, M.
Geiser, D. Hauck, Z. Kraus, A. Phantala & E. Vongphachan (NHMB); 1
spec., Savannakhet prov., 10 E of Savannakhet, Dong Natad forest, 180
m, 4.xii.2011, J. Šťastný leg. (JSCL).
Diagnosis. TL 22–24 mm, elongate, oblong-oval (Fig. 3A).
Head dark green, clypeus yellow. Pronotum and elytra
dark green, with distinct yellow margins laterally; apex of
yellow margin on elytra slightly hooked. Ventral surface
black; with lateral part of metaventrite and metacoxae
yellow; abdominal ventrites III–V with a pale spot on
each side (Fig. 3B). Dorsal surface smooth in both sexes.
Prosternal process lanceolate, with distinct lateral bead,
surface fl at, apex pointed; lateral parts of metaventrite
tongue-shaped, slender, not reaching lateral margins;
metacoxal lines distinct, well impressed, nearly parallel
medially, divergent anteriorly, along metacoxal lines with
a row of punctures. Forelegs reddish-brown, midlegs and
hindlegs reddish-brown to black; protarsomeres 1–3 of
male broadly expanded into a palette, with four rows of
adhesive setae on their ventral side; metatarsal claws single
in both sexes. Median lobe of aedeagus ʻCʼ-shaped in later-
al view, moderately curved (Fig. 3C); gradually narrowed
from base to near apex in ventral view, apex furcate, with
a deep acute-angled notch medially, lateral margins of
apex straight and parallel (Fig. 3D). Parameres moderately
broad, approximately the same width apically (Fig. 3E).
Habitat. In Laos all specimens were collected at lower
altitudes between 180 and 300 m above sea level. The
specimen from Sekong was attracted by a light trap placed
near the border of a lowland river. The single specimen
from Dong Natad was collected in a shallow swamp in low-
land monsoon primary forest, associated with Hyphydrus
lyratus (Swartz, 1808), Hyphydrus sp. and Lacconectus sp.
For habitat photos see H B (2013).
Distribution. China (Shanghai) (Fig. 20A), Indonesia,
Laos, Malaysia, Thailand, Vietnam (R 1899,
Z 1920, F 1936, N 1995, H
& B 2013).
Cybister cf. guerini Aubé, 1838
(Figs 4A–E; 20A)
Cybister guerini Aubé, 1838: 57 (orig. descr.); S (1882): 741 (China,
Manchuria); R (1899): 342 (Guangdong); F (1932): 35
(Guangdong); Z (1989): 91 (Guangdong).
Material examined. CHINA: G: 76 spec., Guangzhou,
14.xi.1964, T. X. Peng got from market (SYSU); 1 1 , Nanhai District,
Xiqiaoshan Mt., 11.x.1973, S. T. Lu leg. (SYSU); 1 2 , Guangzhou
(SYSU). H: 1 , Lingshui County, Diaoluoshan Mt., Xiaomei
Village, 29.xii.1963, L. F. Huang leg. (SYSU).
Diagnosis. TL 31–36 mm, elongate, oblong-oval (Fig. 4A).
Head dark green, clypeus yellow. Pronotum and elytra dark
green to black, with distinct yellow margins laterally; apex
of yellow margin on elytra hooked. Ventral surface black;
abdominal ventrites III–V with a yellow spot on each side
(Fig. 4B). Dorsal surface dimorphic: smooth sometimes
with inconspicuous small granules in male, in female with
isolated short, irregular striae on pronotum, and with dense
longitudinal striae of various lengths on elytra except at its
apex. Prosternal process lanceolate, with distinct lateral
bead, surface fl at, apex pointed; lateral parts of metaven-
trite tongue-shaped, slender, not reaching lateral margins;
metacoxal lines distinct, well impressed, nearly parallel
medially, divergent anteriorly, along metacoxal lines with
a row of punctures. Forelegs reddish-brown, midlegs and
hindlegs reddish-brown to black; protarsomeres 1–3 of male
broadly expanded into a palette, with four rows of adhesive
setae on their ventral side; metatarsal claws single in both
sexes. Median lobe of aedeagus ʻCʼ-shaped in lateral view,
moderately curved (Fig. 4C); slightly narrow near apex in
ventral view, apex furcate, with a notch medially, lateral
margins of apex sinuate (Fig. 4D). Parameres moderately
broad, strap-like apically (Fig. 4E).
Comments to classifi cation. As already mentioned in
H B (2013), it will be necessary to
study the type material of C. guerini and C. limbatus
carefully before identifying freshly collected material. In
many collections, historical specimens of the same species
can be found under the two diff erent names.
Habitat. The knowledge of habitat of Cybister guerini is
very poor, all specimens examined here were collected in
1960s and 70s without habitat information. Some speci-
mens from Guangzhou were obtained from a market.
Distribution. China (Guangdong, Hainan, Hong Kong,
“Manchuria”) (Fig. 20A), India, Indonesia, Laos, Thailand,
Vietnam (R 1899; Z 1920; F
1932, 1933b; G 1938; V 1969; Z
1989; N 1995, N & H 2022b). The record
of C. guerini from Northeast China by S (1882) is
outside the distribution area of this Oriental species and
needs to be confi rmed. First record from Hainan.
Cybister laevis Falkenström, 1936
(Figs 5A–C; 20A)
Cybister laevis Falkenström, 1936: 243 (orig. descr.).
Cybister tripunctatus var. laevis: G (1938: 64).
Type material. H: (NHRS), labelled: “Giufu-Shan [Chong-
qing, Jinfoshan Mt.] / Szechuan / Em. Reitter [p] // Cybister / laevis /
Falk. [hw] / det. Falkenström [p] // Typus [red label with black frame, p]
// 7533 / E91 + [blue label, p] // NHRS-JLKB / 000073974 [p]”.
Diagnosis. See Cybister tripunctatus lateralis. Ventral
side yellowish.
Comments to classifi cation. The female type specimen
has smooth elytra without any striolae – a condition which
can be observed also in all females of C. tripuncatus
lateralis. Since it is an odd, old museum specimen, it
has a yellowish ventral side, which is not the case for C.
tripunctatus lateralis from China, where the ventral side is
always black or ferruginous in freshly hatched specimens.
The size (26 mm) and the coloration of the dorsal side also
indicate that C. laevis is conspecifi c with the variable and
widespread C. tripunctatus lateralis, as already suggested
by G (1938).
Distribution. China (Chongqing) (Fig. 20A).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 81
Fig. 4. Cybister guerini Aubé, 1838 (, Guangzhou, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus
in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Cybister lateralimarginalis torquatus
(Fischer von Waldheim, 1829)
(Figs 6A–E; 19B; 20B)
Dytiscus torquatus Fischer von Waldheim, 1829: 25 (orig. descr.).
Cybister chaudoirii Hochhuth, 1846: 213 (orig. descr.), synonymy by
N (2001: 93).
Cybister hedini Zaitzev, 1908: 419 (orig. descr.), synonymy by N
(2001: 93).
Cybister kansou Feng, 1936: 13 (orig. descr.), synonymy by J et al.
(2010: 257).
Cybister lateralimarginalis chaudoirii: Z (1972): 359 (Xinjiang).
Material examined. CHINA: N: 1 , Yinchuan, 9.–15.ix.1984,
W. Ma leg. (SYSU). X: 1 , Ili River, 28.vii.2005, L. Zhao leg.
(SYSU); 1 1 , Shule County, 21.vii.2006, L. Zhao leg. (SYSU).
W : 42 spec., donated by Xing-min Wang, purchased
from market, without data (SYSU). TAJIKISTAN: 1 1 , Leninabad
[Chudschand], 5.v.1978, Dr. Sobotka leg. (NMPC); 1 , Leninabad,
viii.1983, E. Tamplon leg. (LHCM). UZBEKISTAN: 2 , Tashkent/
Sirdarya Province, 50 km WSW Tashkent, near Chinaz (Yangi Chinoz
Qorgoni), 40°57′N 68°43′E, 9.viii.2004, L. & E. Hendrich leg. (Loc. 5b)
(LHCM); 1 , Tashkent, Bektemir, Chirchiq fl oodplains, 41°15′57.64″N
69°22′58.37″E, 450 m, 21.iv.2009, L. & E. Hendrich leg. (LHCM);
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
82
1 , Tashkent “Botanicheskiy sad”, ponds, 23.–28.vi.2005, L. & E.
Hendrich leg. (Loc. 4) (LHCM); 2 6 , Buchara, Kopon 15 km
Ecocentr Dzejran, 21.v.1992, A. Snizek leg. (LHCM).
Diagnosis. TL 29–37 mm, elongate, oblong-oval (Fig.
6A). Head dark green, clypeus yellow. Pronotum and
elytra dark green to black, with distinct yellow margin
laterally. Ventral surface yellow; middle of metaventrite
and metacoxae reddish-brown to brown (Fig. 6B). Dorsal
surface dimorphic: smooth sometimes with inconspicuous
small granules in male, in female with isolated short, irreg-
ular striae on pronotum, and with dense longitudinal striae
of various lengths on elytra except at its apex. Prosternal
Fig. 5. Cybister laevis Falkenström, 1936 (holotype ). A – habitus in dorsal view; B – habitus in ventral view; C – labels. Scale bar: 5.0 mm.
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 83
process lanceolate, with distinct lateral bead, surface fl at,
apex pointed; lateral parts of metaventrite tongue-shaped,
slender, not reaching lateral margins; metacoxal lines dis-
tinct, well impressed, nearly parallel medially, divergent
anteriorly, along metacoxal lines with a row of punctures.
Forelegs yellow, midlegs and hindlegs yellow to brown;
protarsomeres 1–3 of male broadly expanded into a pal-
ette, with four rows of adhesive setae on their ventral
side. Metatarsal claw single in both sexes. Median lobe of
aedeagus ʻCʼ-shaped in lateral view, strongly curved (Fig.
6C); with acuminate protuberance on each side near apex
in ventral view, apex rounded, slightly infl ated (Fig. 6D).
Parameres moderately broad, strap-like apically (Fig. 6E).
Comments to classifi cation. At fi rst sight, many Central
Asian and Chinese specimens of Cybister lateralimargina-
lis torquatus have a more elongate (i.e. less broadly oval)
Fig. 6. Cybister lateralimarginalis torquatus (Fischer von Waldheim, 1829) (, Shule County, Xinjiang). A – habitus in dorsal view; B – habitus in ventral
view; C – median lobe of aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
84
habitus than European specimens of the nominotypical
subspecies. Furthermore, in lateral view, some specimens
of C. lateralimarginalis ssp. torquatus are slightly more
fl attened than C. lateralimarginalis from Europe. The late-
ral yellow margin on the elytra is wider in some specimens
of C. lateralimarginalis torquatus than in most specimens
of the nominotypical form. The extension of the striolation
of the elytra in females can vary in both forms as does the
shape of the outer posterior angle of the metafemora. The
more individuals we have examined, the clearer it becomes
that Cybister lateralimarginalis is a very variable species
over its area of distribution, and the expression of the cha-
racteristics previously mentioned in the literature for sepa-
rating subspecies can vary greatly (see also, e.g., H et
al. 2019). Further genetic and morphological investigations
are needed to determine whether the current subspecifi c
concept of C. lateralimarginalis is justifi ed or not.
Habitat. There is little knowledge about the habitat of
Cybister lateralimarginalis torquatus in China. Some
specimens were purchased from a market. In Uzbekistan
most specimens were collected from exposed eutrophic
ponds and ditches, mainly with bottle traps. The specimens
from Bukhara were collected near steppe lakes at light.
Distribution. China (Gansu, Ningxia, Nei Mongol, Xin-
jiang) (Fig. 20B), Afghanistan, Georgia, India, Russia,
Tajikistan, Turkmenistan, Turkey, Uzbekistan (Z
1972, G & N 2012, N & H 2022b).
First record from Tajikistan and Uzbekistan.
Fig. 7. Cybister lewisianus Sharp, 1873 (, Guangzhou, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus
in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 85
Cybister lewisianus Sharp, 1873
(Figs 7A–E; 21A)
Cybister lewisianus Sharp, 1873: 46 (orig. descr.); S (1882): 732
(Shanghai); R (1899): 350 (Shanghai, Jiangxi); F
(1932): 36 (Anhui: Wuhu, Beijing, Hubei: Wuchang, Jiangsu: Suzhou,
Zhejiang: Hangzhou and Jiaxing); Z (1981): 110 (Fujian); Z
(1989): 92 (Anhui, Beijing, Hubei, Jiangsu, Zhejiang); L (1992):
36 (Liaoning).
Material examined. CHINA: G: 2 1 , Nanhai Dis-
trict, Jiujiang Township, 16.xi.1957, D. X. Gu leg. (SYSU); 6 2 ,
Guangzhou, 14.xi.1964, T. X. Peng got from market (SYSU). H:
1 , Mt. Wuzhishan at Wuzhishan city, 15–24.vii.2011, Jingke Li leg.
(SYSU). H: 2 2 , Wuchang, vii.1956, Q. Y. Hu leg. (SYSU).
Diagnosis. TL 23–27 mm, elongate, oblong-oval (Fig.
7A). Head dark green, clypeus reddish-brown. Pronotum
and elytra dark green to black, with distinct yellow mar-
gins laterally. Ventral surface black, with lateral part of
metaventrite and metacoxae reddish-brown; abdominal
ventrites III–V with a yellow spot on each side (Fig.
7B). Dorsal surface smooth in both sexes. Prosternal
process lanceolate, with distinct lateral bead, surface
fl at, apex pointed; lateral parts of metaventrite tongue-
shaped, slender, not reaching lateral margins; metacoxal
lines distinct, well impressed, nearly parallel medially,
divergent anteriorly, along metacoxal lines with a row of
punctures. Forelegs yellow, midlegs and hindlegs yellow
to reddish-brown; protarsomeres 1–3 of male broadly
expanded into a palette, with four rows of adhesive setae
on their ventral side; both sexes with a single metatarsal
claw. Median lobe of aedeagus ʻCʼ-shaped in lateral view,
moderately curved (Fig. 7C); slender in ventral view,
apex rounded (Fig. 7D). Parameres moderately broad,
strap-like apically (Fig. 7E).
Habitat. This species inhabits large ponds with aquatic
plants (M & K 1993).
Distribution. China (Anhui, Beijing, Guangdong, Hai-
nan, Hebei, Hubei, Jiangsu, Jiangxi, Liaoning, Shanghai,
Zhejiang) (Fig. 21A), India, Indonesia, Japan, Vietnam
(R 1899; Z 1920; F 1932,
1933b; G 1938; Z 1972; Z
1989; M & K 1993; N 1995; G &
N 2012; N & H 2022b). First record
from Guangdong and Hainan.
Cybister cf. limbatus (Fabricius, 1775)
(Figs 8A–E; 18F; 21A)
Dytiscus limbatus Fabricius, 1775: 230 (orig. descr.).
Dytiscus aciculatus Herbst, 1784: 123 (orig. descr.), synonymy by
S (1808: 10).
Cybister limbatus: S (1882: 739) (Manchuria); Z
(1919: 77) (Taiwan); K (1931: 176) (Taiwan); M & K
(1993: 144) (Taiwan).
Material examined. CHINA: G: 1 , Canton, Honam
Island, P’an-yu District. 25.vii.1934, William E. Hoff mann leg. (SYSU);
1 , Lung-tau Shan, Yiu vill., above Taokokwan, 600 m, 7.vi.1947, PM,
Gressitt & Lam (SYSU); 13 3 , Guangzhou, 14.xi.1964, T. X.
Peng got from market (SYSU). G: 1 , Nanning, 19.vi.1977,
Z. H. Huang leg. (SYSU). H: 1 , Sanya, 28.xii.1963, T. X. Peng
leg. (SYSU); 1 , Jianfengling Mt., 1.xi.1983, P. L. Chen leg. (SYSU);
1 , Limushan Mt., 28.v.1984, G. R. Lin leg. (SYSU). Y: 1 ,
Longling County, 1060 m, 24.vi.1956, B. S. Zhou leg. (SYSU).
Diagnosis. TL 33–39 mm, elongate, oblong-oval (Fig.
8A). Head dark green, clypeus yellow. Pronotum and
elytra dark green to black, with distinct yellow mar-
gins laterally, apex of yellow margin on elytra hooked.
Ventral surface black; abdominal ventrites III–V with a
reddish-brown spot on each side (Fig. 8B). Dorsal sur-
face dimorphic: smooth sometimes with inconspicuous
small granules in male, in female with isolated short,
irregular striae on pronotum and dense longitudinal striae
of various length on elytra except at its apex. Prosternal
process lanceolate, with distinct lateral bead, surface
fl at, apex pointed; lateral parts of metaventrite tongue-
shaped, slender, not reaching lateral margins; metacoxal
lines distinct, well impressed, nearly parallel medially,
divergent anteriorly, along metacoxal lines with a row of
punctures. Forelegs reddish-brown, midlegs and hind-
legs reddish-brown to black; protarsomeres 1–3 of male
broadly expanded into a palette, with four rows of adhe-
sive setae on their ventral side; both sexes with a single
metatarsal claw. Median lobe of aedeagus ʻCʼ-shaped
in lateral view, moderately curved (Fig. 8C); gradually
narrowed from base to near apex in ventral view, apex
furcate, with a deep obtuse-angled notch medially, lateral
margins of apex concave (Fig. 8D). Parameres moderately
broad, strap-like apically (Fig. 8E).
Comments to classifi cation. As mentioned under C.
guerini, it will be necessary to study the type material
of C. limbatus and C. guerini to clarify the status of
both taxa. Specimens of the same morphospecies can be
found under the two diff erent names in many historical
collections (H B 2013).
Habitat. This species inhabits deep ponds and abandoned
paddy fi elds with aquatic plants (M & K
1993, H B 2013); specimens from
Danxiashan Mt. (Guangdong) were found in a temporary
pond in a valley (Fig. 18F). In Thailand, the adults prey
on larvae and pupae of Aedes aegypti (Linnaeus, 1762)
and Culex quinquefasciatus Say, 1823 (Y 2005).
The species is sold as food in Thailand (C et al. 1998)
and Laos (H B 2013).
Distribution. China (Guangdong, Guangxi, Hainan,
Taiwan, Xizang, Yunnan, “Manchuria”) (Fig. 21A),
Afghanistan, India, Japan, Laos, Myanmar, Nepal, Paki-
stan, Philippines, Thailand, Vietnam (R 1899;
Z 1920; F 1932, 1933b; G
1938; V 1969; Z 1972; M & K
1993; N 1995; G & N 2012; H
& B 2013; N & H 2022b). The
record of C. limbatus from Northeast China by S
(1882) is outside the distribution area of this Oriental
species and needs to be confi rmed. First records from
Guangdong, Guangxi and Hainan.
Cybister rugosus (W. S. Macleay, 1825)
(Figs 9A–E; 18D–E, G; 19C; 21A)
Dytiscus rugosus W. S. Macleay, 1825: 32 (orig. descr.).
Cybister indicus Aubé, 1838: 62 (orig. descr.), synonymy by S
(1882: 992).
Cybister rugosus: N (1995): 78 (Taiwan).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
86
Material examined. CHINA: C: 1 , Fengdu County,
Shiping Village, 610 m, 5.x.1994, J. Chen leg. (SYSU); 2 , Fengdu
County, Shiping Village, 580 m, 6.x.1994, J. Yao leg. (SYSU). G-
: 1 2 , suburbs of Guangzhou, 6.xi.1956 (SYSU); 1 , Nanhai
District, Jiujiang Township, 16.xi.1957, D. X. Gu leg. (SYSU); 69 spec.,
Guangzhou, 14.xi.1964, T. X. Peng got from market (SYSU); 1 , Sans-
hui District, 18.xi.1964, T. X. Peng leg. (SYSU); 2 , Nanhai District,
Xiqiaoshan Mt., 11.x.1973, S. T. Lu leg. (SYSU); 1 , Guangzhou, Sun
Yat-sen University, x.1973, S. T. Lu leg. (SYSU); 1 , Dinghushan Mt.,
28.vi.1978, Z. Y. Wang leg. (SYSU); 1 , Guangzhou, Sun Yat-sen
University, in front of Huaishitang, 2.x.1999, L. Z. Hua leg. (SYSU); 1
, Shaoguan, Danxiashan Mt., 30.v.–8.vi.2008, F. L. Jia leg. (SYSU); 1
, Fengkai County, He’erkou, light trap, 2.xii.2010, L. J. Yang & Y. L.
Yu leg. (SYSU). H: 1 , Hwang-mei & vicinity, Hwang-mei Dist.,
9.–11.vii.1935, H. W. Djou (SYSU). H: 1 , Tongdao County,
20.viii.1982, Z. H. Huang leg. (SYSU); 1 , Hengyang, Zhoujiaao,
28.vi.1986, Q. Zou leg. (SYSU). J: 1 , main peak of Jinggangshan
Mt., 1.ix.2011, L. J. Yang leg. (SYSU); 1 , Nanchang (SYSU); 1 ,
Shusi, H. Y. Cai leg. (SYSU). Y: 1 , Yingjiang County, 820 m,
25.v.1983, L. Z. Hua leg. (SYSU). Z: 2 , Kecheng district,
Fig. 8. Cybister limbatus (Fabricius, 1775) (, Guangzhou, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of
aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 87
Lankeshan Mt., Quzhou, 28.8781N, 118.9199E, 118 m, 18.iii.2020, Z.
Y. Jiang leg. (SYSU).
Diagnosis. TL 27–33 mm, elongate, oblong-oval (Fig.
9A). Head dark green, clypeus yellow. Pronotum and
elytra dark green to black, with distinct yellow margins
laterally, apex of yellow margin on the elytra hooked.
Ventral surface black with lateral part of metaventrite and
metacoxae yellow; abdominal ventrites I–V with yellow
spot on each side (Fig. 9B). Dorsal surface dimorphic:
smooth sometimes with inconspicuous small granules
in male; in female with isolated short, irregular striae on
pronotum and dense longitudinal striae of various length
on elytra at its apex. Prosternal process lanceolate, with
distinct lateral bead, surface fl at, apex pointed; lateral parts
of metaventrite tongue-shaped, slender, not reaching lateral
margins; metacoxal lines distinct, well impressed, nearly
parallel medially, divergent anteriorly, along metacoxal
lines with a row of punctures. Forelegs yellow, midlegs and
Fig. 9. Cybister rugosus (W. S. Macleay, 1825) (, Fengkai County, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median
lobe of aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
88
hindlegs yellow to black; protarsomeres 1–3 of male broad-
ly expanded into a palette, with four rows of adhesive setae
on their ventral side; both sexes with a single metatarsal
claw. Median lobe of aedeagus ʻCʼ-shaped in lateral view,
moderately curved (Fig. 9C); slightly narrow near apex in
ventral view, apex furcate, with a notch medially, lateral
margins of apex concave (Fig. 9D). Parameres moderately
broad, strap-like apically (Fig. 9E).
Habitat. This species inhabits ponds, abandoned paddy
fi elds and wetlands with aquatic plants (M & K-
1993); specimens from Shenzhen (Guangdong) were
found in a lowland marsh (Figs 18D–E) and specimens
from Lankeshan Mt. (Zhejiang) were found in a natural
pond (Fig. 18G).
Distribution. China (Chongqing, Guangdong, Hubei,
Hunan, Jiangxi, Taiwan, Yunnan, Zhejiang) (Fig. 21A),
Cambodia, Indonesia (Java, Kalimantan), Japan, Laos,
Malaysia, Singapore, Thailand, Vietnam (R
1899, Z 1920, G 1938, M
& K 1993, N 1995, H et al. 2004,
H & B 2013). First record from
Chongqing,
Guangdong, Hubei, Hunan, Jiangxi, Yun-
nan and Zhejiang.
Cybister tripunctatus lateralis (Fabricius, 1798)
(Figs 10A–E; 18D–F; 19D; 21B)
Dytiscus lateralis Fabricius, 1798: 64 (orig. descr.).
Cybister asiaticus Sharp, 1882: 731 (orig. descr.), synonymy by N
(2001: 90).
Cybister gotschii Hochhuth, 1846: 214 (orig. descr.), synonymy by
N (2001: 90).
Cybister similis Régimbart, 1899: 352 (orig. descr.), synonymy by
N (2001: 90).
Cybister tripunctatus orientalis Gschwendtner, 1931: 99 (orig. descr.),
synonymy by N (2001: 90); F (1932): 37 and F (1933a):
329 (Sichuan, Hubei: Wuchang, Fujian: Fuzhou and Xiamen, Guang-
dong, Hainan); B (1946): 151 (Heilongjiang); Z (1981):
110 (Fujian); Z (1989): 91 (Fujian, Guangdong, Hainan, Hubei,
Hunan, Jiangsu, Sichuan, Yunnan, Xizang, Zhejiang); L (1992):
36 ( NE China); P et al. (1992): 484 (Yunnan); M & K
(1993): 142 (Taiwan).
Cybister szechwanensis Falkenström, 1936: 238 (orig. descr.), synonymy
by J et al. (2010: 258).
Cybister tripunctatus lateralis: J et al. (2010): 258 (NE China, N China,
Fujian, Guangdong, Hainan, Hubei, Hunan, Jiangsu, Sichuan, Taiwan,
Yunnan, Xizang, Zhejiang).
Material examined. CHINA: C: 1 , Jinyunshan Mt., 800
m, 13.vi.1994, Y. W. Zhang leg. (SYSU). F: 1 , Haitong, NR.,
Foot How, iii.1926, H. T. Chen (SYSU); 1 , Fuqing, Xitai, 6.–13.x.1932
(SYSU); 1 , Nan-ping District, Yen-ping, vi.–vii.1933, D. C. Ngu
(SYSU). G: 1 2 , Ch’eng-hai District, K’ei-luk, Swa-
tow, viii.1932, Y. C. Ng (SYSU); 1 , Lingnan University, 10.vi.1948
(SYSU); 3 , suburbs of Guangzhou, 6.xi.1956 (SYSU); 1 2 ,
Guangzhou, Honam Island, 2.–9.xi.1957, D. X. Gu leg. (SYSU); 6
6 , Nanhai District, Jiujiang Township, 16.xi.1957, D. X. Gu leg.
(SYSU); 1 2 , Guangzhou, Xinshi, 11.x.1964, C. M. Chen & Z.
W. Huang leg. (SYSU); 1 , Guangzhou, Lianhe, 18.x.1964, Z. Y. Chen
leg. (SYSU); 1 , Guangzhou, Baiyunshan Mt., 2.xi.1964, J. C. Bao leg.
(SYSU); 126 spec., Guangzhou, 14.xi.1964, T. X. Peng got from market
(SYSU); 1 , Guangzhou, 18.xi.1964, T. X. Peng leg. (SYSU); 1 , Nan-
hai District, Shijing Village, 10.x.1973, S. T. Lu leg. (SYSU); 84 spec.,
Nanhai District, Xiqiaoshan Mt., 10.–11.x.1973, S. T. Lu leg. (SYSU); 1
2 , suburbs of Guangzhou, Renhe Township, 26.–27.x.1973, S. T.
Lu leg. (SYSU); 1 , Nanhai District, 2.xi.1973, S. T. Lu leg. (SYSU); 1
, Shunde District, Xingtan Township, 2.xi.1973, S. T. Lu leg. (SYSU);
1 , Guangzhou, Baiyunshan Mt., 6.xi.1973, S. T. Lu leg. (SYSU); 1 ,
Ruyuan County, Luoyang Commune, 26.v.1974, C. Liao leg. (SYSU);
1 , Guangzhou, Kangle, 28.iv.1989, F. L. Jia leg. (SYSU); 1 1 ,
Shenzhen, Noi Ling Ding Island, 20.x.1998, H. D. Chen leg. (SYSU);
1 1 , Xinhui District, viii.2001, X. L. Tong leg. (SYSU); 4 11
, Gaoming District, Yangmei Township, 20.–27.iv.2006, F. L. Jia et
al. leg. (SYSU); 1 , Lian County, Dadongshan Mt., 3.–9.vii.2008, W.
C. Xie leg. (SYSU); 1 , Lianjiang City, 26. ix.2008 (SYSU); 1 , Feng-
kai County, Heishiding, 5.x.2011, Y. Li leg. (SYSU). G: 2 ,
Rong County, 10.x.1962, D. X. Gu leg. (SYSU); 4 2 , Nanning,
19.vi.1977, Z. H. Huang leg. (SYSU). H: 1 , Jianfengling Mt.,
18.xii.1963, T. X. Peng leg. (SYSU); 3 1, Sanya, 24.xii.1963, T.
X. Peng leg. (SYSU); 1 , Dongfang-Huangliu, 25.xii.1963, T. X. Peng
leg. (SYSU); 2 , Tongshi, 27.xii.1963, T. X. Peng leg. (SYSU); 1
, Lingshui County, Diaoluoshan Mt., Xiaomei Village, 29.xii.1963,
L. F. Huang leg. (SYSU); 1 , Qionghai, Shangyong Village, light
trap, 17.vii.1984, C. L. Wang leg. (SYSU). H: 1 , Zigui County,
Maoping Township, 80 m, 28.iv.1994, W. Z. Li leg. (SYSU). H:
2 , Tongdao County, 20.viii.1982, Z. H. Huang leg. (SYSU); 1 ,
Hengyang, Zhoujiaao, 2.vii.1986, Q. Zou leg. (SYSU). J: 1 ,
Nanchang, iii.1963, X. W. Chen leg. (SYSU); 1 , Nanchang (SYSU).
S: 1 , Shanghai, 16.ii.1926, W. E. Hoff mann (SYSU). Y:
1 , Xishuangbanna, Menglun Township, light trap, 580 m, 13.ix.1993,
H. L. Xu leg. (SYSU); 1 , Xishuangbanna Tropical Botanical Garden,
650 m, 9.viii.2003, Y. Zhang leg. (SYSU). X: 1 , Gyirong County,
Gyirong Commune, 26.vii.1984, Z. X. Yan leg. (SYSU); 5 10 ,
Linzhi, Mêdog County, Beibeng Township, Didong Village, 840 m,
16.viii.2006, M. Bai leg. (SYSU); 3 1 , Linzhi, Mêdog County,
Beibeng Township, 887 m, 17.viii.2006, M. Bai leg. (SYSU); 3 2
, Linzhi, Mêdog County, Yarang Township, Didong Village, 760 m,
19.viii.2006, M. Bai leg. (SYSU); 20 spec., Linzhi, Mêdog County, 1084
m, 21.viii.2006, M. Bai leg. (SYSU); 20 spec., Linzhi, Mêdog County,
1084 m, 23.viii.2006, M. Bai leg. (SYSU); 1 , Linzhi, Mêdog County
80K, 2110 m, 24.viii.2006, M. Bai leg. (SYSU).
Diagnosis. TL 24–28 mm, elongate, oblong-oval (Fig. 10A).
Head dark green, clypeus yellow. Pronotum and elytra dark
green to black, with distinct yellow margins laterally, apex
of yellow margin on the elytra slightly hooked. Ventral
surface black, sometimes with lateral part of metaventrite
and metacoxae yellow; abdominal ventrites III–V with a
yellow spot on each side (Fig. 10B). Dorsal surface smooth
in both sexes. Prosternal process lanceolate, with distinct
lateral bead, surface fl at, apex pointed; lateral parts of
metaventrite tongue-shaped, slender, not reaching lateral
margins; metacoxal lines distinct, well impressed, nearly
parallel medially, divergent anteriorly, along metacoxal lines
with a row of punctures. Forelegs reddish-brown, midlegs
and hindlegs reddish-brown to black; protarsomeres 1–3
of male broadly expanded into a palette, with four rows of
adhesive setae on their ventral side; metatarsal claws single
in both sexes. Median lobe of aedeagus ʻCʼ-shaped in lateral
view, moderately curved (Fig. 10C); slightly narrow near
apex in ventral view, apex furcate, with a deep acute-angled
notch medially, lateral margins of apex convex (Fig. 10D).
Parameres moderately broad, tapering apically (Fig. 10E).
Habitat. The most common species of the genus in
Southeast Asia. An eurytopic species which is widespread
in artifi cial habitats such as irrigation channels, fl ooded
paddy fi elds, open swamps, fi sh and ornamental ponds and
larger ditches, even in urban and suburban areas (M &
K 1993, H et al. 2004). Specimens from
Shenzhen (Guangdong) were found in a lowland marsh (Figs
18D–E) and specimens from Danxiashan Mt. (Guangdong)
were found in a temporary pond in a valley (Fig. 18F).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 89
Distribution. China ( Chongqing, Fujian, Guangdong,
Guangxi, Hainan, Hebei, Heilongjiang, Hubei, Hunan,
Jiangsu, Jiangxi, Macao, Shanghai, Sichuan, Taiwan,
Yunnan, Xizang, Zhejiang) (Fig. 21B), Afghanistan, United
Arab Emirates, Azerbaijan, Bhutan, Cyprus, India, Iran,
Iraq, Japan, Kyrgyzstan, Laos, Mongolia, Nepal, Oman,
Pakistan, Russia, Singapore, Syria, Thailand, Tajikistan,
Turkmenistan, Turkey, Uzbekistan (R 1899;
Z 1920; F 1932, 1933a,b; G
1938; V 1969; Z 1972; Z 1989; M &
K 1993; H et al. 2004; G & N
2012; H & B 2013; N & H
2022b). First record from Chongqing, Guangxi, Jiangxi
and Shanghai.
Fig. 10. Cybister tripunctatus lateralis (Fabricius, 1798) (, Guangzhou, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median
lobe of aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
90
Cybister ventralis Sharp, 1882
(Figs 11A–E; 22A)
Cybister ventralis Sharp, 1882: 742 (orig. descr.); K (1931): 177
(Taiwan); F (1933a): 329 (Sichuan, Shandong: Qingdao).
Material examined. None from China. INDIA: 1 2 , Behar, India
or., Coll. A. Zimmermann (ZSMG). SRI LANKA: 1 , Negombo, 1899,
W. Horn, Coll. A. Zimmermann (ZSMG).
Diagnosis. TL 28–32 mm, broad, oblong-oval (Fig. 11A).
Head dark green, clypeus yellow. Pronotum dark green
to black with lateral yellow margins projecting a little
inwards along anterior margin behind eyes; elytra dark
green to black, with distinct yellow margin laterally, apex
of yellow margin on elytra hooked. Ventral surface yellow
with central part of metaventrite and metacoxae black;
abdominal ventrites III–V yellowish with a dark brown
band anteriorly (Fig. 11B). Dorsal surface dimorphic in
females: smooth or with isolated short, irregular striae on
pronotum and dense longitudinal striae of three-fourths
of elytral length from base. Prosternal process lanceolate,
with distinct lateral bead, surface fl at, apex pointed;
lateral parts of metaventrite tongue-shaped, slender, not
reaching lateral margins; metacoxal lines distinct, well
impressed, nearly parallel medially, divergent anteriorly,
along metacoxal lines with a row of punctures. Forelegs
and midlegs yellow, tarsi darker; metafemur and metatibia
yellow with broad dark brown marking; female with a
Fig. 11. Cybister ventralis Sharp, 1882 (, Pusa, Bihar, India). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus
in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 91
single metatarsal claw. Median lobe of aedeagus ʻCʼ-
shaped in lateral view, moderately curved (Fig. 11C);
slightly narrow near apex in ventral view, apex furcate,
with a notch medially, lateral margins of apex concave
(Fig. 11D). Parameres more elongate, strap-like apically
(Fig. 11E).
Distribution. China (Macao, Sichuan, Shandong, Taiwan)
(Fig. 22A), Bangladesh, India, Myanmar, Pakistan, Sri
Lanka (R 1899, Z 1920, K 1931,
F 1933a, V 1969, N 1995, G &
N 2012, N & H 2022b).
Subgenus Melanectes Brinck, 1945
Type species. Cybister owas Laporte, 1835.
Diagnosis. Without distinctive yellow or pale margins on
the elytron and pronotum. Males with a single metatarsal
claw; females with two claws, the posterior claw small.
Included (Chinese) species. Cybister brevis Aubé, 1838;
C. convexus Sharp, 1882; C. danxiaensis sp. nov.; C. pos-
ticus Aubé, 1838; C. sugillatus Erichson, 1834.
Cybister brevis Aubé, 1838
(Figs 12A–E; 18G; 22A)
Cybister brevis Aubé, 1838: 98 (orig. descr.); K (1931): 177 (Taiwan);
F (1936): 238 (Sichuan); L (1992): 36 (Liaoning).
Material examined. CHINA: G: 1 , Jiangkou County,
Shengxisi, 19.vi.1984 (SYSU); 1 , west slope of Fanjingshan Mt.,
Huguosi, 1.viii.2001, H. Pang leg. (SYSU). H: 1 , Longshan
County, Tongche, 5.vi.1965, Z. Y. Chen leg. (SYSU). Z: 2
1 , Quzhou, Kecheng district, Lankeshan Mt., 28.8781N, 118.9199E,
118 m, 18.iii.2020, Z. Y. Jiang leg. (SYSU).
Fig. 12. Cybister brevis Aubé, 1838 (, Lankeshan Mt., Zhejiang). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus
in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
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Diagnosis. TL 20–25 mm, elongate, oblong-oval (Fig.
12A). Head dark green, clypeus orange. Pronotum and el-
ytra dark green to black, apex of elytra with reddish-brown
spots. Ventral surface black; abdominal ventrites III–IV
with a yellow spot on each side (Fig. 12B). Head and prono-
tum densely covered with coarse punctures, elytra sparsely
covered with inconspicuous small granules. Pro sternal
process lanceolate, with distinct lateral bead, surface
fl at, apex pointed; lateral parts of metaventrite tongue-
shaped, slender, not reaching lateral margins; metacoxal
lines distinct, well impressed, nearly parallel medially,
divergent anteriorly, along metacoxal lines with a row of
punctures. Forelegs reddish-brown, midlegs and hindlegs
reddish-brown to black; protarsomeres 1–3 of male broadly
expanded into a palette, with four rows of adhesive setae
on their ventral side; male with a single metatarsal claw;
female with two metatarsal claws, posterior claw small
and sinuate. Median lobe of aedeagus ʻCʼ-shaped in lateral
view, slightly curved, gradually broader from base to apex,
then strongly narrowing at apex (Fig. 12C); broad medially
in ventral view, apex obtuse-angled (Fig. 12D). Parameres
moderately broad, strap-like apically (Fig. 12E).
Habitat. This species inhabits ponds, abandoned paddy
fi elds and ditches of paddy fi elds with aquatic plants (M
Fig. 13. Cybister convexus Sharp, 1882 (, Kunming, Yunnan). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus in
lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 93
& K 1993); specimens from Lankeshan Mt. (Zhe-
jiang) were found in a natural pond (Fig. 18G).
Distribution. China (Guizhou, Hunan, Liaoning, Si-
chuan, Taiwan, Zhejiang) (Fig. 22A), Japan, South Korea
(R 1899, Z 1920, F 1933a,
G 1938, Z 1972, M & K
1993, N 1995, N & H 2022b). First
record from Guizhou, Hunan and Zhejiang.
Cybister convexus Sharp, 1882
(Figs 13A–E; 18H; 19E; 22A)
Cybister convexus Sharp, 1882: 718 (orig. descr.); R (1899):
353 (Yunnan); F (1932): 35 (Yunnan); V (1969): 284
(Yunnan); Z (1989): 93 (Yunnan).
Material examined. CHINA: Y: 1 , Tia Feng on (Grassy hill 3ml.
N Yunnanfu). 25.viii.1934, C. Brownell leg. (SYSU); 1 , Honghe Hani-
Yi Autonomous Prefecture, Lvchun County, Niukong Township Section,
terrace, 22.9873N, 102.2675E, 1336 m, 30.iv.2021, Z. Y. Jiang, Z. M. Yang,
B. P. Huang & Z. Q. Mai leg. (SYSU); 1 , Dehong Dai Aut. Pref. mount.
Range 31 km E Luxi, 2280 m, 24°29′31″N 98°52′58″E, 3.vi.2007, D.W.
Wrase leg. (LHCM); 1 , Yunnan, Baoshan Pref. mount. Range, 22 km S
Tengchong, 1750m, 24°49′29″N 98°29′27″E (in fi shpond), 30.viii.2009,
D. W. Wrase leg. (LHCM).
Diagnosis. TL 24–30 mm, elongate, oblong-oval (Fig. 13A).
Head, pronotum and elytra dark green to black, apex of elytra
with reddish-brown spots. Ventral surface black; abdominal
ventrites III–IV with a yellow spot on each side (Fig. 13B).
Head and pronotum smooth, elytra densely covered with
conspicuous small granules. Prosternal process lanceolate,
with distinct lateral bead, surface fl at, apex pointed; lateral
parts of metaventrite tongue-shaped, slender, not reaching
lateral margins; metacoxal lines distinct, well impressed,
nearly parallel medially, divergent anteriorly, along metacox-
al lines with a row of punctures. Legs black; protarsomeres
1–3 of male broadly expanded into a palette, with four rows
of adhesive setae on their ventral side; male with a single
metatarsal claw; female with two metatarsal claws, posterior
claw small and sinuate. Median lobe of aedeagus ʻCʼ-shaped
in lateral view, slightly curved, broadest at middle, strongly
narrowing in apical fi fth, apex rounded (Fig. 13C); broad
medially in ventral view, apical fi fth slender like a tubular
spine, apex truncated (Fig. 13D). Parameres moderately
broad, strap-like apically (Fig. 13E).
Habitat. The habitat information is not well known for this
species. It can be found from hilly to high mountain regions
(A et al. 2018). In Yunnan, it was collected in a
water-fi lled rut on a forest track from Dehong (H
2008) and in a fi sh pond. In Lvchun County it was collected
at night from terraced fi elds (Fig. 18H).
Distribution. China (Yunnan) (Fig. 22A), India, Thailand
(R 1899; Z 1920; F 1932, 1933b;
G 1938; V 1969; Z 1989; G
& N 2012; A et al. 2018).
Cybister (Melanectes) danxiaensis sp. nov.
(Figs 14A–E; 15A–E; 22B)
Type locality. China, Guangdong Province, Shaoguan, Renhua County,
Danxiashan Mt., ca. 25.0244N, 113.7158E.
Type material. H: (SYSU), labelled: “广东 仁化 丹霞 / 1941.
VIII / 林平 [hw] // CHINA: Guangdong Province / Renhua County /
Danxiashan Mt. / viii.1941, Ping Lin leg. [p] // HOLOTYPE / CYBISTER
/ danxiaensis sp. nov. / Jiang, Zhao, Mai & Jia det. 2021 [red label, p]”.
Description. Habitus (Fig. 15A) elongate oblong oval,
with continuous outline, widest in apical third of elytra,
moderately convex. Dorsal surface distinctly matt. TL 21.7
mm; TL-h 19.4 mm. MW 11.3 mm.
Colouration. Head black; pronotum black with broad
brown sides; elytra blackish brown; appendages black;
ventral side blackish brown.
Head moderately broad, ca. 0.6× width of pronotum,
trapezoidal. Surface densely covered with fi ne punctures.
A pair of depressions present near anterior margin of cly-
peus. Anterior margin of clypeus concave. Labium and
genae with some punctures near gular suture. Antenna with
antennomeres long and slender.
Pronotum transverse (width/length ratio = 2.36), broa-
dest between posterior angles, lateral margins moderately
curved, without lateral bead. Punctation double; coarse
setigerous puncture row present along anterior margin, with
a perpendicular setigerous puncture row laterally that does
not reach posterior margin; fi ne punctures dense throughout
whole surface. Centre of disc with a moderately impressed
longitudinal line medially.
Elytra with dense large protuberances that are arranged
almost in rows; each protuberance with a puncture medially
in which is a small granule (Fig. 14A). Punctation double;
larger punctures sparse between protuberances; finer
punctures dense throughout whole surface. Medioapical
area of elytra with a fi eld of short, coarse setae (Fig. 14B).
Ventral side (Fig. 15B). Prosternal process lanceolate,
with distinct lateral bead, surface fl at, apex pointed (Fig.
Fig. 14. Morphological characters of Cybister danxiaensis sp. nov. A –
detail of elytral structure; B – Medioapical area of elytra viewed with
sutural margin at bottom of picture; C – prosternal process; D – metacoxal
lines; E – mesotaromeres. Scale bars: 1.0 mm.
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
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14C). Lateral parts of metaventrite tongue-shaped, slender,
not reaching lateral margins; metacoxal lines distinct, well
impressed, nearly parallel medially, divergent anteriorly,
along metacoxal lines with a row of punctures (Fig. 14D).
Lateral part of abdominal ventrites without spot.
Legs. Protarsomeres 1–3 with four rows of adhesive
setae on their ventral side, broadly expanded into a palette;
claws simple. Mesotibia with setae in a continuous line on
posteroapical margin; mesotarsomeres with two separate
series of setae present along posteroventral and posterodor-
sal apical margin; claws simple (Fig. 14E). Hindlegs lost.
Male genitalia. Median lobe of aedeagus ʻCʼ-shaped in
lateral view, strongly curved, broadest at middle, strongly
narrowing in apical fourth, apex rounded (Fig. 15C); in
ventral view broad at base, apical fourth slender like a
tubular spine, apex rounded (Fig. 15D). Parameres mo-
derately broad, distinctly curved in apical third, strap-like
apically (Fig. 15E).
Female. Unknown.
Diff erential diagnosis. Based on the pronotum and elytra
without distinct lateral yellow margins, the new species
can be assigned to subgenus Melanectes (M et al.
Fig. 15. Cybister danxiaensis sp. nov. (holotype ). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus in lateral view;
D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 95
2007). Within the subgenus, the new species is similar to
Cybister sugillatus but can easily be recognized by the large
punctate protuberances on the elytra and the longer apical
slender part of the median lobe of the aedeagus (elytra
smooth without punctate protuberances, apical slender
part of median lobe of aedeagus shorter in C. sugillatus).
Etymology. The species is named after the type locality,
Danxiashan Mt., Guangdong Province, China; the name
is an adjective.
Distribution. So far known only from the type locality in
northern Guangdong, China (Fig. 22B).
Cybister posticus Aubé, 1838
(Figs 16A–E; 22B)
Cybister posticus Aubé, 1838: 87 (orig. descr.); J et al. (2010): 259
(Xizang: Zayü County).
Material examined. CHINA: X: 1 , Mêdog County, Beibeng
Township, 800 m, 5. ix.1974, F. S. Huang leg. (SYSU); 1 , Bomê
County, Yigong, 1450 m, 9.v.1983, Y. H. Han leg. (SYSU); 1 1 , Zayü
County, Xia Zayü, 28.50N 97.00E, 4.–8.x.2007, F. M. Shi leg. (SYSU).
Diagnosis. TL 23–35 mm, elongate, oblong-oval (Fig. 16A).
Head dark green, clypeus reddish-brown. Pronotum and
elytra dark green to black, apex of elytra with reddish-brown
Fig. 16. Cybister posticus Aubé, 1838 (, Zayü County, Xizang). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of aedeagus
in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
96
spots. Ventral surface black; abdominal ventrites III–IV with
a yellow spot on each side (Fig. 16B). Head and pronotum
smooth, elytra densely covered with conspicuous small
granules. Prosternal process lanceolate, with distinct lateral
bead, surface fl at, apex pointed; lateral parts of metaven-
trite tongue-shaped, slender, not reaching lateral margins;
metacoxal lines distinct, well impressed, nearly parallel me-
dially, divergent anteriorly, along metacoxal lines with a row
of punctures. Forelegs reddish-brown, midlegs and hindlegs
reddish-brown to brown; protarsomeres 1–3 of male broadly
expanded into a palette, with four rows of adhesive setae
on their ventral side; male with a single metatarsal claw;
female with two metatarsal claws, posterior claw small and
sinuate. Median lobe of aedeagus ʻCʼ-shaped in lateral view,
slightly curved, broadest at middle, strongly narrowing in
apical fi fth, apex rounded (Fig. 16C); broad medially in
ventral view, apical fi fth slender like a tubular spine, apex
rounded (Fig. 16D). Parameres moderately broad, strap-like
apically (Fig. 16E).
Habitat. Although this species is widely distributed in
India, nothing is known about its habitat.
Distribution. China (Xizang) (Fig. 22B), Bangladesh, In-
Fig. 17. Cybister sugillatus Erichson, 1834 (, Shenzhen, Guangdong). A – habitus in dorsal view; B – habitus in ventral view; C – median lobe of
aedeagus in lateral view; D – median lobe of aedeagus in ventral view; E – paramere. Scale bars: 5.0 mm (A, B), 1.0 mm (C–E).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 97
dia, Nepal, Pakistan (R 1899, Z 1920,
V 1969, J et al. 2010, G & N 2012).
Cybister sugillatus Erichson, 1834
(Figs 17A–E; 18D–E, H; 19F; 22B)
Cybister sugillatus Erichson, 1834: 227 (orig. descr.); R (1899):
355 (Xizang); Z (1919): 77 (Taiwan); K (1931): 176
(Taiwan); F (1932): 36 (Beijing, Fujian: Fuzhou and Xiamen,
Guangdong, Hainan, Hubei: Wuchang, Jiangxi, Sichuan, Xizang,
Zhejiang); F (1936): 238 (Sichuan); Z (1981): 110
(Fujian); Z (1989): 92 (Beijing, Fujian, Guangdong, Hainan,
Hubei, Jiangxi, Sichuan, Xizang, Zhejiang); M & K
(1993): 142 (Taiwan).
Cybister bisignatus Aubé, 1838: 88 (orig. descr.), synonymy by S
(1882: 991).
Cybister notasicus Aubé, 1838: 90 (orig. descr.), synonymy by S
(1882: 717).
Cybister olivaceus Boheman, 1858: 21 (orig. descr.), synonymy by
S (1882: 717).
Material examined. CHINA: G: 8 7 , Guangzhou,
14.xi.1964, T. X. Peng purchased from market (SYSU); 1 , Ruyuan
County, Tianjingshan Mt., 8.viii.1980, Q. X. Zhang leg. (SYSU); 1
, Fengkai County, Heishiding, 10.iv.1985, W. Wu leg. (SYSU); 1 ,
Fengkai County, Heishiding, 14.vii.1999, Y. C. Wang leg. (SYSU); 1
, Fengkai County, Heishiding, 12.vii.2000, Y. L. Feng leg. (SYSU);
2 2 , Gaoming District, Yangmei Township, 20–27.iv.2006,
D. P. Wu, N. Yu, Q. S. Liu & Y. Y. Niu leg. (SYSU); 1 , Zhuhai,
back mountain of Sun Yat-Sen University, 5.–8.vii.2011, F. L. Jia leg.
(SYSU). G: 1 , Rong’an County, Dajiang-Yayao, 28.iv.1959
(SYSU); 2 1 , Nanning, iii.2001, L. Hojný leg. (NMPC); 1 ,
Jingxi, Bangliang Nature Reserve, 1.viii.2010, J. H. Huang leg. (SYSU).
H: 1 , Cheung-kon-ts’uen, Ka-luk-kong (18ml. E. of Nam-fung),
Kiung-shan Dist., 29.–31.iii.1935, F. K. To leg. (SYSU); 1 , Lingshui
County, 1.i.1964, T. X. Peng leg. (SYSU); 1 , Jiangfengling, the third
partition, 24.xi.1981, J. P. Lu leg. (SYSU); 1 , Jiangfengling, Heiling,
10.vii.1982, L. Z. Hua leg. (SYSU). H K: 1 , Hong Kong,
1934 (SYSU). Y: 1 1 , Chengjiang County, 30.iv.1932, Z.
L. Pu leg. (SYSU).
Diagnosis. TL 18–23 mm, elongate, oblong-oval (Fig.
17A). Head, pronotum and elytra dark green to black,
lateral part of pronotum reddish-brown, apex of elytra
with reddish-brown spots. Ventral surface black; abdomi-
nal ventrites III–IV with a yellow spot on each side (Fig.
17B). Dorsal surface smooth. Prosternal process lanceo-
late, with distinct lateral bead, surface fl at, apex pointed;
lateral parts of metaventrite tongue-shaped, slender, not
reaching lateral margins; metacoxal lines distinct, well
impressed, nearly parallel medially, divergent anteriorly,
along metacoxal lines with a row of punctures. Forelegs
reddish-brown, midlegs and hindlegs reddish-brown to
black; protarsomeres 1–3 of male broadly expanded into
a palette, with four rows of adhesive setae on their ventral
side; male with a single metatarsal claw; female with two
metatarsal claws, posterior claw small and sinuate. Medi-
an lobe of aedeagus ʻCʼ-shaped in lateral view, strongly
curved, broadest at middle, strongly narrowing in apical
fi fth, apex rounded (Fig. 17C); broad medially in ventral
view, apical fi fth slender like a tubular spine, apex rounded
(Fig. 17D). Parameres moderately broad, strap-like apically
(Fig. 17E).
Habitat. This species inhabits not only ponds, wetlands,
abandoned paddy fi elds and ditches of paddy fi elds with
aquatic plants, but also eutrophic waters near grazing land
(M & K 1993). Specimens from Shenzhen
(Guang dong) were found in in a lowland marsh (Figs
18D–E) and specimens from Lvchun County (Yunnan)
were found in terraced fi elds (Fig. 18H).
Distribution. China (Beijing, Fujian, Guangdong,
Guangxi, Hainan, Hebei, Hong Kong, Hubei, Jiangxi,
Sichuan, Taiwan, Xizang, Yunnan, Zhejiang) (Fig. 22B),
Afghanistan, Bhutan, India, Indonesia, Japan, Laos, Malay-
sia, Myanmar, Nepal, Pakistan, Philippines, Singapore, Sri
Lanka (R 1899; Z 1920; F 1932,
1933b; G 1938; V 1969; Z
1972; Z 1989; M & K 1993; N 1995;
H et al. 2004; G & N 2012; H
& B 2013; N & H 2022b). First
record from Guangxi and Yunnan.
Key to the Chinese species of Cybister
1 Pronotum and elytra with yellow margins laterally;
female with a single metatarsal claw. Subgenus Cy-
bister Curtis, 1827 ....................................................2
– Pronotum and elytra without yellow margins late-
rally; female with two metatarsal claws. Subgenus
Melanectes Brinck, 1945 ....................................... 11
2 Apex of median lobe of aedeagus furcate in ventral
view. ....................................................................... 3
– Apex of median lobe of aedeagus not furcate in ven-
tral view. ................................................................. 9
3 Metaventrite and metacoxae black with lateral parts
yellow. .................................................................... 4
– Metaventrite and metacoxae entirely black. ........... 7
4 Smaller species, TL: 22–28 mm; female with dorsal
surface smooth (Fig. 3A). ....................................... 5
– Larger species, TL: 27–33 mm; female dorsal surface
with dense longitudinal striae of various lengths on
elytra except at its apex (Fig. 11A). ....................... 6
5 Smaller species, TL: 22–24 mm; median lobe of ae-
deagus with apical notch shallower, lateral margins
of apex straight and parallel in ventral view (Fig.
3D). .................................... C. fumatus Sharp, 1882
– Larger species, TL: 24–28 mm; median lobe of ae-
deagus with apical notch deeper, lateral margins of
apex convex in ventral view (Fig. 10D). ...................
C. tripunctatus lateralis (Fabricius, 1798) (partim)
and C. laevis Falkenström, 1936
6 Abdominal ventrites III–V black, with lateral yellow
spots (Figs 1B; 9B). ...... C. rugosus (W. S. Macleay,
1825) and C. bengalensis Aubé, 1838
– Abdominal ventrites III–V yellowish, with a dark
brown band anteriorly (Fig. 11B). .............................
........................................... C. ventralis Sharp, 1882
7 Smaller species, TL 24–28 mm; median lobe of aedea-
gus with acute-angled apico-medial notch (Fig. 10D).
..... C. tripunctatus lateralis (Fabricius, 1798) (partim)
– Larger species, TL>30.0 mm; median lobe of aedea-
gus with an obtuse-angled apico-medial notch. ..... 8
8 Median lobe of aedeagus in ventral view with apex
and base almost equal in width, lateral margins of
apex sinuate, notch of apex shallower (Fig. 4D). ......
.............................................. C. guerini Aubé, 1838
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
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Fig. 18. Habitats of Cybister in China. A–C – lake, Shanxi (C. chinensis); D–E – lowland marsh, Shenzhen, Guangdong (C. rugosus, C. tripunctatus
lateralis and C. sugillatus); F – temporary pond in a valley, Danxiashan Mt., Guangdong (C. tripunctatus lateralis and C. limbatus); G – natural pond,
Lankeshan Mt., Zhejiang (C. rugosus and C. brevis); H – terraced fi elds pools, Lvchun County, Yunnan (C. convexus and C. sugillatus).
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 99
– Median lobe of aedeagus in ventral view with apex
narrower than base, lateral margins of apex concave,
notch of apex deeper (Fig. 8D). ................................
................................... C. limbatus (Fabricius, 1775)
9 Smaller species, TL: 23–27 mm; ventral surface
black with lateral parts of metaventrite and meta-
coxae reddish-brown (Fig. 7B); median lobe of
aedeagus slender in ventral view (Fig. 7D); female
with dorsal surface smooth. .....................................
........................................ C. lewisianus Sharp, 1873
– Larger species, TL>29 mm; ventral surface yellow
with middle parts of metaventrite and metacoxae
reddish-brown or dark green; median lobe of aedea-
gus thick in ventral view; female dorsal surface with
isolated short, irregular striae on pronotum, and with
dense longitudinal striae of various lengths on elytra
except at its apex. ................................................. 10
10 Larger species, TL: 35–40 mm; prosternal process
and middle parts of metaventrite and metacoxae dark
green (Fig. 2B); median lobe of aedeagus in ventral
view ridged in middle, lateral margin strongly curved
near apex, apex obtuse-angled (Fig. 2D). .................
............................... C. chinensis Motschulsky, 1854
– Smaller species, TL: 29–37 mm; prosternal process
yellow, middle parts of metaventrite and metacoxae
reddish-brown (Fig. 6B); median lobe of aedeagus in
ventral view with acuminate protuberance on each side
near apex, apex rounded, slightly infl ated (Fig. 6D). ...
................................ C. lateralimarginalis torquatus
(Fischer von Waldheim, 1829)
Fig. 19. Live specimens of Chinese Cybister. A – C. chinensis Motschulsky, 1854 (mating couple); B – C. lateralimarginalis torquatus (Fischer von
Waldheim, 1829); C – C. rugosus (W. S. Macleay, 1825); D – C. tripunctatus lateralis (Fabricius, 1798); E – C. convexus Sharp, 1882; F – C. sugillatus
Erichson, 1834.
JIANG et al.: Review of Cybister of China (Coleoptera: Dytiscidae)
100
Fig. 20. Map of distribution of Cybister in China. Fig. 21. Map of distribution of Cybister in China.
Fig. 22. Map of distribution of Cybister in China.
11 Elytra with large protuberances densely arranged al-
most in rows, each protuberance with a medial pun-
cture containing a small granule (Fig. 15A). .............
............................................. C. danxiaensis sp. nov.
– Elytra smooth or with small granules. .................. 12
12 Head and pronotum densely covered with coarse
punctures (Fig. 12A); apical fi fth of median lobe
broad (Figs 12C–D). ............... C. brevis Aubé, 1838
– Head and pronotum smooth; apical fi fth of median
lobe of aedeagus slender, like a tubular spine. ..... 13
13 Elytra smooth in both sexes (Fig. 17A). ...................
..................................... C. sugillatus Erichson, 1834
– Elytra with small granules in both sexes. ............. 14
14 Median lobe of aedeagus thicker in ventral view,
apex truncated (Fig. 13D). .......................................
.......................................... C. convexus Sharp, 1882
– Median lobe of aedeagus thinner in ventral view,
apex rounded (Fig. 16D). ... C. posticus Aubé, 1838
Discussion
The unique specimen of Cybister (Melanectes) danxia-
ensis sp. nov. was collected from Danxiashan Mountain
eighty years ago. Around twenty years ago we examined
and dissected the specimen and considered it as represen-
ting an undescribed species. We would have liked to have
a few more specimens available for the description, and
therefore went to the type locality several times – unfor-
tunately without success. Most probably, the species’
Acta Entomologica Musei Nationalis Pragae, volume 63, number 1, 2023 101
former habitat at the locus typicus has been destroyed
due to construction of the river channel or to agricultural
development. No other specimens of this species were
collected in well-preserved forests near the type locality
either. Danxiashan Mountain has a very special landform
clearly diff erent from that of nearby areas.
Although Cybister includes the largest predatory wa-
ter beetles in China, and members of several species are
occasionally sold at local food markets, their distribution
and habitat requirements are still poorly known. Further
investigations, using more eff ective collecting methods
such as baited bottle traps (e.g., A R 1985,
H 1987, V et al. 2013, H et al.
2019), could increase the knowledge of the species’ ecolo-
gical demands and their actual conservation status in China.
Acknowledgements
We are grateful to Miss Fang Chen and Zai-xiong Chen,
the managers of Danxiashan National Nature Reserve. We
thank Dr. Xing-min Wang, a specialist on ladybird beetles
in South China Agricultural University (Guangzhou, Chi-
na), for his donation to some specimens of C. chinensis and
C. lateralimarginalis torquatus. Michael Balke (ZSMG)
is thanked for providing some photos of median lobes and
parameres; and Johannes Bergsten (NHRS), for providing
the photos of an important type specimen. Martin Spies
(ZSMG) is thanked linguistic corrections. This study
was supported by GDAS Special Project of Science and
Technology Development (2020GDSYL-20200102021,
2020GDASYL- 20200301003).
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