2008

Asiatic Herpetological Research, Vol. 11

pp. 17–23

On the Status of the Chinese Pitviper Ceratrimeresurus shenlii Liang and
Liu in Liang, 2003 (Serpentes, Viperidae), with the Addition ofProtobothrops cornutus (Smith, 1930) to the Chinese Snake Fauna
PATRICK DAVID1,*, HAIYAN TONG2, GERNOT VOGEL3 AND MINGYI TIAN4
1

Département Systématique et Evolution, USM 602 Taxonomie-collection – Reptiles et Amphibiens,
CP 30, Muséum National d’Histoire Naturelle, 57 rue Cuvier, F-75231 Paris Cedex 05, France,
2
16, cour du Liégat, F-75013 Paris, France,
3
Im Sand 3, D-69115 Heidelberg, Germany,4
College of Natural Resources & Environment, South China Agricultural University, Wushan,
Guangzhou 510640, People’s Republic of China.
*Corresponding author E-mail: pdavid@mnhn.fr

Abstract.- Ceratrimeresurus shenlii Liang and Liu in Liang, 2003 was described as a new genus and a new specieson the basis of the first “horned” specimen of pitviper recorded from the People’s Republic of China. This taxon has
been overlooked in the literature. The original description is here translated verbatim into English. The holotype iscompared with other horned pitvipers known from Asia. On the basis of its scalation and pattern, Ceratrimeresurus
shenlii is synonymized with Protobothrops cornutus (Smith, 1930). The range of this latter species, previously
endemic to Vietnam, is expanded northeastwards by approximately 780 airline km. A brief comment on the zoogeography of South China is given.
Keywords.- Serpentes, Ceratrimeresurus, Ceratrimeresurus shenlii, Protobothrops cornutus, China, Vietnam, taxonomy, zoogeography.

Introduction
A new Chinese genus and new species of Asian pitviper,
Ceratrimeresurus shenlii Liang and Liu in Liang, 2003
has remained overlooked in the literature.
Ceratrimeresurus shenlii was not included in the latest
checklist of the Trimeresurus complex (Gumprecht etal., 2004), nor announced in Wolfgang Wüster’s invaluable website “Venomous Snakes Systematic Alert”(http://biology.bangor.ac.uk/~bss166/update.htm).
Lastly, this taxon was not considered by Malhotra andThorpe (2004, 2005) in their revision on the pitvipers of
the Trimeresurus complex.
The description of Ceratrimeresurus shenlii Liangand Liu in Liang (2003: 411; Plate 8: Fig. 13. Type locality: “Working site 02 at Wuzhishan forest, Ruyuan
Xian”, Guangdong Province) appeared in a chapter of abook on the natural history of Nanling Nature Reserve,
located in Nanling Mountains, in the north of
Guangdong Province (Pang, 2003), a fact that may
explain that the new taxon remained overlooked by theherpetological community. However, this new species
was merely mentioned in September 2004 in the forumof a website dedicated to venomous snakes(http://www.venomdoc.com/forums - last viewed on© 2008 by Asiatic Herpetological Research

May 30th, 2005), where this species was regarded, without explanations, as a synonym of Protobothrops cornutus. It was also tentatively regarded as a synonym ofProtobothrops cornutus by Vogel (2006).
Only three other taxa of Asian pitvipers with moreor less raised supraoculars were previously known fromthe mainland: Trimeresurus wiroti (see David et al.,2006), Protobothrops cornutus and Triceratolepidophis
sieversorum. In this paper, we provide a translation of
the original description. The new taxon is, according to
the re-examination of the holotype, compared with thecurrently known horned species of mainland Asia and its
status is discussed.

Materials and Methods
Body and tail lengths were measured to the nearest mm.
The number of ventral scales is counted according to
Dowling (1951). The terminal scute is not included in
the number of subcaudals. The numbers of dorsal scalerows are given at one head length behind head, at midbody (i.e. at the level of the ventral plate corresponding
to a half of the total number of ventrals), and at one headlength before vent, respectively. Values for symmetrichead characters are given in left/right order.

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Abbreviations of measurements and other meristic
characters.MEASUREMENTSAND
RATIOS:
HL:
head
length; SVL: snout-vent length; TaL: taillength; TL: total length; TaL/TL: ratio tail length /total
length.
MERISTIC CHARACTERS: DSR: formula of dorsal scalerows; MSR: number of dorsal scale rows at midbody;
IL: infralabials; SC: subcaudals; SL: supralabials; VEN:
ventrals.
Museum abbreviations.- BMNH: Natural History
Museum, London, UK. - FMNH: Field Museum of
Natural History, Chicago, USA. - MNHN: Muséum
National d’Histoire Naturelle, Paris, France. ZFMK: Zoologisches Forschungsmuseum Alexander
Koenig, Bonn, Germany.

Results
Translation of the original description ofCeratrimeresurus shenlii.- The original description
appeared in Liang (2003: 411), but is credited to Liangand Liu. It was published in Chinese with a short
English summary. A poor quality, small color photograph of the holotype appears on Plate 8, Figure 13 of
the book. A verbatim translation of the original description should read as follows (numbers placed in square
brackets refer to our comments placed after this translation):[1]
Jiao laotietou Ceratrimeresurus shenlii Liang[2]
and Liu, gen. and sp. nov. (See colour plate 13 )The upper eyelid forms on each side a raised triangle
covered with small scales, the base of the triangle beingslightly like a three-sided pyramid; the length of the triangular horn is 1.5 times the diameter of the eye, and theheight from the tip of the triangle to the edge of theupper eyelid is equal to the eye diameter. The tip of thesnout is blunt, the upper snout surface is broad and flattened, with the lateral edge slightly upturned. The lengthof the snout is about a 1/4 of head length and 1/3 of headwidth. The nostril is placed near the tip of the snout,
slightly directed sideward and almost rounded in shape,
the opening being in the middle of a divided nasal scale;there is one upper nasal scale. There are 5 small scales[3]
between the two nostrils . The upper head surface iscovered with granular scales, which become progressively larger and more keeled from the front to the backof the head. The cephalic surface is slightly convex andbecomes progressively flat and broader backwards.
[4]
There are 7 small interorbital scales . There are 11
small scales surrounding the base of the triangular horn.
The tip of the triangular horn is one rather large scale, to2008

which is adjacent a smaller scale which makes the horn
looking as bifurcated. The eye is rather large and thethird of its volume is out of the head. The iris is a vertically elongated oval. There is a row of small scales
around the orbit and many scales between the snout sideand temporal region. The forward sunken part of theocciput is roughly equal to the length of the snout, withthe end of the neck plunged in it. The neck is slender, its
diameter being less than half of the head width. There[5]
are 14 supralabial scales , among which the first andsecond ones are separated from the nasal scales by a rowof 3 small scales, and the third to fifth ones do not enter[6]
the loreal pit , but are separated from it by a row ofsmall scales; the edge of the upper jaw is convex downwards; 14 infralabials, the fourth being the largest, thefirst to the fourth ones in contact with the anterior chinshields, and the posterior chin shields being rather large.
One large fang is located on the anterior part of theupper jaw, followed by 2 smaller teeth. No teeth on the[7]palatine and pterygoid . Five small teeth are present on
the lower jaw. The loreal pit is placed antero-inferiorly
to the eye.
The head is peach-shaped and slightly flattened,with a narrow and elongated neck; the body is rather
wide but slender and long, with a long, pointed tail. The
dorsal scales are strongly keeled, 23-23-21-17 rows, therows of cervical scales are more oblique than those of
the body scales. There are 187 ventral scales +77 pairsof subcaudals. The anal scale is single. The vertebral
scales are normal.
The back of the body is of a grey color, whichbecomes light grey on the sides. On the head upper surface, there is a ‘X’ shaped pattern, made of a blotch
extending from the nasal scale to the front of the opposite triangular, and of a blackish brown elongated blotch
extending from the posterior of the horn through the
rearward supraorbital scale (behind the horn) backwards
up to the posterior of the head. A white streak extends
from the posterior margin of the eye through the temporal up to the posterior of the head, followed by a blackish
brown stripe from the corner of the mouth to the sides of
the throat. The upper and lower lips are marked withblackish brown square shaped spots. Diffuse and irregular spots cover the rest of the body throughout. The dorsal side of the head and the body is brown in color.
Dorsal blotches extend on each side from the middle ofthe body to the vent, the right and the left rows being offset and these blotches being linked together by theirinner corner; they are merged together in the middle ofthe body and linked each other again by the inner corneron the posterior part of the body, then merged together
again from the vent to the end of the tail. Light greybrown spots on the lower part on the sides of the body,roughly square in shape or almost rounded, oval or2008

Asiatic Herpetological Research, Vol. 11

reduced to short crossbars. The venter is pale grayishbrown, without other spot in the middle, so that the ventral side of the body is light grey-brown color throughout. There is a pointed scale on the tip of the tail. The
total length of the specimen: 362 + 78 = 440 mm, lengthof the tail/total length = 0.564, weight = 24 g.
The specimen was collected in the thatch on the topof a house near a forest, coiled with the head in the middle of its body. Not very active. It was knocked downwith a little stick on the ground and captured. The locality is the “Working site 02” at Wuzhishan forest, Ruyuan[8]Xian , July 1996. The specimen is deposited in the
Laboratory of Zoology of the Faculty of Biotechnology,[9]
Jinan University .
Comments on the original description. - In this originaldescription, no mention of the sex of the specimen is
given, although it is a female according to the shape ofthe base of the tail. Other comments are:[1]: Translation: “Horned iron-head [snake]”. The vernacular name “Laotietou” is given in China to
Protobothrops mucrosquamatus, due to the shapeof its head, looking like an ancient iron.
[2]: It should rather appear as Plate 8: fig. 13.
[3]: This value includes the two supranasals plus thescales separating these latter ones.
[4]: If one counts the cephalic scales on a line connecting the middle of the supraoculars, the value is 14(see below).
[5]: This value is obviously erroneous; we counted 8 SL.
[6]: This is obviously a lapsus for the orbit.
[7]: this lack of teeth on the palatine bones is surprisingand may be an artifact. Teeth are very commonly
miscounted, as they are masked by the tissues andone usually needs to peel back the gum tissues tocount the sockets. All the tooth counts in thisdescription may be unreliable (A. Malhotra, pers.
comm, January 2006).
[8]: Ruyuan County, in northern Guangdong Province,close to the Guangdong-Hunan border. The
Nanling National Nature Reserve is located in thecentre of Nanling Mountains, which extend from
northeast Guangxi to southwest Fujian.
[9]: Guangzhou, Guangdong Province.
Redescription of the holotype of Ceratrimeresurus
shenlii (Figs. 1-3).- The original description includes
some imprecision or mistakes. In combining data of theoriginal description and our own data, the morphology is
as follows:
Body moderately stout; head subtriangular, wide at its
base, clearly distinct from the neck, thick and swollen
when seen from the side, depressed between the uplifted19

areas of the supraoculars. Snout average in relative
length, about one quarter of HL, bluntly rounded when
seen from above, depressed in its center, obliquely truncated when seen from the side, with a distinct canthus
rostralis. Eye relatively large. Tail long and tapering.
The holotype is now in average condition and somewhat
dessicated:
SVL: 362 mm; TaL: 78 mm; TL: 440 mm; ratio
TaL/TL: 0.177.
VEN: 187; SC: 77, paired, plus one terminal scale; anal
shield entire.
DSR: 23-21-17 scales, rhomboid, distinctly keeled.
Rostral visible from above, broader than high, triangular; nasals subrectangular, divided, with a round nostril opening near the tip of the snout, directed slightly
sideward and almost rounded in shape; 2 internasals oneach side, separated by 2 small scales; 4/4 canthal
scales, slightly larger than adjacent snout scales, bordering the canthus rostralis; 2 elongate upper preoculars
above the loreal pit; lower preocular forming the lower
margin of loreal pit; 3/3 small postoculars; 5/5 supraoculars on an uplifted triangular area, of which the twocentral supraocular scales are strongly enlarged, triangular and strongly obliquely erected (“horn like”) andextending out of the head margin, 1.5 times the diameterof the eye, convergent and originating from the samebase covered with small scales; 8 or 9 slightly enlarged
scales on upper snout surface on a line between thescales separating the internasals and a line connecting
the anterior margins of eyes, smooth, juxtaposed, irregular in shape; 14 cephalic scales on a line between thebase of the supraoculars (including the scales covering
the uplifted basal area), smooth, flat and juxtaposed;
occipital scales larger than cephalic scales, distinctly
keeled; temporal scales small, obtusely keeled; 9 SL, 1st
SL separated from nasal; 2nd SL bordering the anterior
margin of the loreal pit, 3rd SL largest, separated fromthe subocular by 1 scale; 4th SL much smaller than 3rd
SL, separated from the subocular by 1 scale; 5th and posterior SL much smaller; 14 IL, first four pairs in contact
with anterior chin shields.
The coloration and pattern agree with that of theoriginal description. The light postocular streak is quite
conspicuous, but the overall pattern of the body is rather
faded.

Discussion
Nomenclatural considerations.- The combined description of the genus and species was brief and may raisesome questions about its validity. Two points need to be
discussed. The first one relates to the combined description of the genus Ceratrimeresurus and of its sole20

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2008

Figure 1. Ceratrimeresurus shenlii, holotype. Generalview of body. Photograph by Tian Mingyi.

Figure 2. Ceratrimeresurus shenlii, holotype. Generalview of body. Photograph by Tian Mingyi.

Figure 3. Ceratrimeresurus shenlii, holotype. Lateralview of the head. Photograph by Tian Mingyi.

Figure 4. Head of the second Chinese specimen ofProtobothrops cornutus. Photograph by Fu Jie.

included species, shenlii. According to Art. 13.4 of theInternational Code of Zoological Nomenclature (1999),
in the present case, such a “description of a new nominal
genus and a single included new nominal species isdeemed to confer availability on each name under
Article 13.1.1.” Art. 13.1.1 requires that every new namepublished be accompanied by a description purported todifferentiate the taxon. This combined description ishence considered to be complying with the requirementsof the Code.
The second point relates to the identification of theholotype. No number was cited in the description, andthis specimen was still deposited in Prof. Liang Qishen’s
private collection in December 2005. Nevertheless, as
the name of the collection into which the specimen will
be eventually deposited was clearly indicated in theoriginal description, and as it was confirmed to us thatthe specimen will eventually be deposited in this collection, we consider that this point complies with Art.
16.4.2 of the Code and makes valid the description of

the specific nomen.
A comparison of Ceratrimeresurus shenlii with otherhorned species.- In the Asian mainland, only
Protobothrops cornutus and Triceratolepidophis sieversorum have erected supraoculars. Trimeresurus wiroti
has only slightly raised supraoculars (see David et al.,2006). Another species occurring in western Indonesia,
Trimeresurus brongersmai, has erected horn-likesupraoculars. However, this latter species, related to
Trimeresurus wiroti, differs from the other horned
species by several characters, including the shape of its
snout (David et al., 2006). On the basis of Ziegler et al.
(2001), Herrmann et al. (2002), Herrmann and Ziegler(2002), Herrmann et al. (2004) and of specimens examined by ourselves, a comparison between horned species
and Ceratrimeresurus shenlii is given in Table 1.
Triceratolepidophis sieversorum differs by its
greater number of ventral scales and a different structureof horns, free and strongly divergent. This species also

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pattern is also similar, especially the dorsal blotches, theupper head dark pattern and the postocular streak.
On the basis of the similarities in morphologicalcharacters, we synonymize Ceratrimeresurus shenlii
Liang and Liu in Liang (2003) with Trimeresurus cornutus Smith, 1930, now Protobothrops cornutus (seeHerrmann et al. [2004] for the generic position of thisspecies).
At the generic level, the point to be now discussed
is if the horned species Trimeresurus cornutus Smith,1930 should be referred to a genus distinct fromProtobothrops, in which all other included species arehornless. In this case, the generic nomen
Ceratrimeresurus is available. Pending molecular analyses that should clarify the relationships between the
Chinese and the Vietnamese populations, we have torely only on morphology. Only two currently recognizedgenera of mainland Asia include species with truly erected supraoculars (“horns”), namely Protobothrops (onlyfor P. cornutus) and Triceratolepidophis (T. sieversorum). On the basis of the similarities between P. cornutus and C. shenlii, we adopt a conservative approach inconsidering that erected supraoculars have been convergently evolved in several lineages. Consequently, we
synonymize the genus Ceratrimeresurus Liang and Liu
in Liang (2003) with Protobothrops Hoge and Romano
Hoge, 1983.
A second Chinese specimen of Protobothrops cornutus has appeared on Internet in summer 2005 (Fig. 4),
from Shimentai Nature Reserve (24° 22′-24° 31′ N, 113°05′-113° 31′ E), Nanling Mountains, Yingde County,
Guangdong Province (see Jim and Xu [2002] for a
description of the area). The characters visible on thispicture of a freshly killed specimen are identical with

Figure 5. Distribution map of of Protobothrops cornutus.

shows a peculiar structure of keels of dorsal scales,
which is not found in other horned pitvipers (Ziegler etal., 2001). The dorsal keels of Ceratrimeresurus shenlii
are narrow and made of a single ridge.
In contrast, Ceratrimeresurus shenlii cannot be distinguished from Protobothrops cornutus otherwise thanby minor characters. In both species, the horns stemfrom the same base and are free only in their outermost
part. All scalation characters are similar, including the
keeling of the dorsal scales (see Table 1). Other characters include the keeled occipital scales, the large 3rd SL
separated from the subocular by 1 scale row, 4th and 5th
separated from the subocular by 1 or 2 scales, the number of internasals and of supraoculars. The sole difference bears on the numbers of pairs of infralabials in contact with the anterior chin shield, first 3 pairs in P. cornutus vs. first 4 pairs in Ceratrimeresurus shenlii. The

Table 1. A comparison between known specimens of Asian horned pitvipers.
Taxon

Sex

Tal/TL

MSR

Ven

Sc

Cep

SL

IL

Sup/OcSupOc“C. shenlii”

F

0.177

21

187

77

14

9/?

14/14

5/5

convergent

P. cornutus (1)

M

0.203

21

189

78

16

9/9

14/14

6/6

convergent

P. cornutus (2)

F

0.184

21

192

71

13

9/9

13/14

6/6

convergent

P. cornutus (3)

F

0.182

21

193

72

13

9/9

12/13

7/7

convergent

T. sieversorum (4)

M

0.167

23

228

82

15

8/9

13/14

2/2

divergent

T. sieversorum (5)

F

0.157

22

235

79

15

8/9

13/14

2/2

divergent

T. sieversorum (6)

?

—

21

—

—

16

10/10

13/14

—

divergent

List of cited specimens. Protobothrops cornutus. (1) ZFMK 75067, Phong Nha-Ke Bang National Park, Quang Binh Province, Vietnam (not seen; after
Herrmann et al., 2004); (2) BMNH 1946.1.19.25, “Fan-si-pan Mts., Tonking”, now Mt. Phang Si Pang, Lai Châu Province, Vietnam; (3) MNHN 1937.35,“Tonkin”, northern Vietnam. - Triceratolepidophis sieversorum. (4) ZFMK 71262 (holotype), Phong Nha village, Phong Nha-Ke Bang Nature Reserve,
Quang Binh Province, Vietnam; (5) ZFMK 75066, Phong Nha-Ke Bang Nature Reserve, Quang Binh Province, Vietnam; (6) FMNH 255258, Hin Nammo
NBCA, Boualapha District, Khammouan Province, Laos (specimens (5) and (6) not seen; after Herrmann et al., 2002).

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Asiatic Herpetological Research, Vol. 11

those of Ceratrimeresurus shenlii. The pattern is muchsimilar to that of Protobothrops cornutus depicted in
Herrmann et al. (2004). According to the data kindly
communicated to us by Mr. Fu Jie, the author of the photograph, this second specimen was seen in May 2005. It
was held in alcohol in the home of a member of the Yao
minority, to be most likely used as a medicinal beverage.
The locality of this second specimen lies in a hilly area
at approximately 75 airline km southeastwards from thetype locality of Ceratrimeresurus shenlii in Nanling
Nature Reserve. According to the Yao owner of thisspecimen, this snake was caught by himself while he
was acting as a guide to a scientific survey of the local
herpetological fauna. Several specimens were collected,
but all died in captivity within some days. This species
is there considered very rare.

Conclusions
The occurrence of Protobothrops cornutus in China
makes a considerable northeastward range extension forthis latter species, previously considered endemic to
Vietnam (Nguyên et al., 2005) (see map on Fig. 5).
Ziegler et al. (2006) recorded a specimen ofProtobothrops cornutus from Ha Giang Province, in
extreme northern Vietnam, close to the border withYunnan and Guangxi Zhuang Autonomous Provinces.
The previously northernmost known locality, Mt. Phang
Si Pang (formerly Mt. Fan Si Pan) and Nanling
Mountains are separated by about 980 airline kilometers. Chinese specimens are separated from the new
locality cited in Ziegler et al. (2006) by about 760 airlinekm.
We also consider that the wide gap between the
Chinese population and its Vietnamese relatives may
most likely reflect a lack of appropriate collecting effort
in elevated areas of Guangxi Zhuang Autonomous andGuangdong Provinces than a real geographical gap.
Protobothrops cornutus should be searched for in forestsof mountain or hill ranges such as Daming Shan andDayao Shan (Guangxi Zhuang Autonomous Province)
and in various hills of northern Guangdong locatedbetween the Nanling Mountains and the mountain
ranges of Guangxi Autonomous Region. The herpetofauna of this latter province is still quite poorly known.
Besides the provinces of Guangxi and Guangdong, thisspecies should be searched for in forested areas of southern Yunnan (see Herrmann et al., 2004). Herrmann et al.
(2004) also showed that, in Central Vietnam, P. cornutus
also occurs in the lowlands. However, lowland areas ofsouthern China are quite dry (Anonymous, 1998). Theselowlands separate the hill or mountain ranges of southern China which share a subtropical humid climate with2008

high annual amounts of rainfall (above 2000 mm), a situation which leads to the isolation of the populations of
P. cornutus in this region.
A discussion on the zoogeographical affinities
between North Vietnam and various regions of South
China are outside the scope of this paper, but a merecomparison between the snake faunas of northern
Vietnam and the hills of Guangxi and Guangdong suggests strong similarities. The occurrence ofProtobothrops cornutus in northern Guangdong, as well
as of Shinisaurus crocodilurus and Amphiesma bitaeniatum in northern Vietnam (David et al., 2005; respectively Le and Ziegler, 2003) reinforces the zoogeographicalrelationships of these areas connected by more or lesscontiguous hill or mountain ranges receiving high annual amounts of rainfall.

Acknowledgments
We are indebted to Q. Liang (Guangzhou, China), who
was instrumental in letting TM re-examine the holotype
of Ceratrimeresurus shenlii. We are grateful to A.
Malhotra (Bangor, United Kingdom), M. W. Lau (HongKong, China) and T. Ziegler (Köln, Germany) for theircomments on the draft of this paper; we especially thank
T. Ziegler who made available and permitted us to cite
his paper in press. Lastly, we thank J. Fu (Guangzhou,
China) and N. Wu (Beijing, China) for their kind permission to use in our paper the picture of the second Chinese
specimen of Protobothrops cornutus.

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Submitted: 13 June 2006
Accepted: 04 April 2007