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A new species of the genus Rhabdophis Fitzinger, 1843 (Squamata: Colubridae)

from Guangdong Province, southern China

Article  in  Zootaxa · February 2014

DOI: 10.11646/zootaxa.3765.5.5 · Source: PubMed

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 Zootaxa 3765 (5): 469–480 ISSN 1175-5326 (print edition)
www.mapress.com /zootaxa /
Copyright © 2014 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3765.5.5
http://zoobank.org/urn:lsid:zoobank.org:pub:0DF6D148-ECEF-4589-826C-16BFEF58260E

A new species of the genus Rhabdophis Fitzinger, 1843 (Squamata: Colubridae)

from Guangdong Province, southern China

GUANG-XIANG ZHU1,2, YING-YONG WANG3,6, HIROHIKO TAKEUCHI4 & ER-MI ZHAO1,5,6

1
Life Science College of Sichuan University, Key Laboratory of Bio-resources and Eco-environment (Ministry of Education), Sichuan
Chengdu 610064, P.R. China
2
College of Life and Basic Sciences, Sichuan Agricultural University, Sichuan Ya'an 625014, P.R. China
3
State Key Laboratory of Biocontrol/The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangdong Guangzhou
510275, P.R. China
4
Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto, 606-8502, Japan
5
Chengdu Institute of Biology, Chinese Academy of Sciences, Sichuan Chengdu 610041, P.R. China
6
Corresponding author. E-mail: wangyy@mail.sysu.edu.cn; zem006@163.com

Abstract

A new species, Rhabdophis guangdongensis sp. nov., is described from the Guangdong Province, China. It can be easily
distinguished from other known congeners by cyt b and c-mos sequences, and by the following combination of morpho-
logical characters: body size small; head distinct from the neck; 20 maxillary teeth, the three most posterior teeth strongly
enlarged, and not separated by diastemata from other teeth; six supralabials, the third and fourth touching the eye; seven
infralabials, the first four in contact with anterior chin shields; dorsal scales in 15 rows throughout the body, weakly
keeled, the outer row smooth; 126 ventrals; 39 paired subcaudals; anal scale divided; 44 pairs of narrow dorsolateral black
cross-bars on body and 15 pairs on tail; body and tail with two dorsolateral longitudinal brownish-red lines, respectively
with a series of white spots in cross-bars. The description of this new species brings the total number of described species
of this genus to 21 and represents the tenth known Rhabdophis species in China.

Key words: Colubridae, morphology, mitochondrial and nuclear DNA, taxonomy

Introduction

The genus Rhabdophis Fitzinger (1843) was previously referred to the genus Natrix sensu lato. In the middle of the
20th century, the genus Rhabdophis was revised and diagnosed primarily based on the combination of the
following characters: hemipenes and sulcus spermaticus divided; last two maxillary teeth strongly enlarged,
recurved and usually preceded by a diastema; terrestrial; internasals broad anteriorly, nostrils lateral; apical pits
present or absent; vertebral glands present in several species (Malnate 1960).
Currently, the genus Rhabdophis, containing 20 species, is distributed across Eastern and South Asia (Günther
1858, 1864; Boulenger 1893, 1900; Wall 1923; Bourret 1935; Smith 1943; Malnate 1960; Taylor 1966; Jiang &
Zhao 1983; Malnate & Underwood 1988; Zhao & Adler 1993; Zhao 1997; Stuebing & Tan 2002; de Lang & Vogel
2006; Takeuchi 2013). Nine of these species occur in China, i.e., R. tigrinus (Boie, 1826); R. subminiatus
(Schlegel, 1837); R. nigrocinctus (Blyth, 1855); R. himalayanus (Günther, 1864); R. swinhonis (Günther, 1868); R.
nuchalis (Boulenger, 1891); R. leonardi (Wall, 1923); R. pentasupralabialis (Jiang & Zhao, 1983) and R. adleri
(Zhao, 1997).
During our field surveys in Southern China from 2008 to 2012, we collected an unidentified species of the
genus Rhabdophis that can be externally distinguished from all known congeners based on morphological
characters. In addition, phylogenetic relationship based on the mitochondrial cytochrome b (cyt b) and nuclear
oocyte maturation factor (c-mos) gene sequences revealed that this taxon is differentiated from other congeners in
China. Therefore, we refer this specimen to be a new species which is described in this study.

Accepted by Z. T. Nagy: 10 Jan. 2014; published: 20 Feb. 2014 469

Material and methods

Taxa sampling and data collection. A specimen of Rhabdophis guangdongensis sp. nov. was collected from the
Aizhai Village, Renhua County, Guangdong Province, China. The fresh liver tissues were taken and immediately
preserved in 95% ethanol. Then, the specimen was fixed in 10% buffered formalin and later transferred to 70%
ethanol. In addition, 14 other samples of the genus Rhabdophis, including R. adleri, R. himalayanus, R. leonardi, R.
nigrocinctus, R. nuchalis, R. pentasupralabialis, R. subminiatus, R. tigrinus and R. swinhonis that represent all nine
currently recognized species in China, were used for molecular analyses in this study. Table 1 lists all the samples
used in this study.

TABLE 1. Information of all specimens used in the study.

ID Current genus and species Locality Voucher GenBank accession numbers
name specimen
c-mos cyt b
1 Natrix natrix Sheppey Island, Kent, United LSUMZ 41506 AF471121 AF471059
Kingdom
2 Amphiesma stolatum Bago Div., Myanmar CAS206560 AF471097 AF471030
3 Rhabdophis himalayanus Kachin State, Myanmar CAS224420 KF800919 KF800929
4 Rhabdophis swinhonis Taiwan KUZR 18977 AB861888 AB842176
5 Rhabdophis guangdongensis Aizhai, Renhuay, Guangdong, SYS r000018 KF800920 KF800930
sp. nov. China
6 Rhabdophis adleri Hainan, China WJC20090801 KF800921 KF800931
7 Rhabdophis leonardi Dali, Yunnan, China RDQ 200905367 KF800922 KF800932
8 Rhabdophis leonardi Panzhihua, Sichuan, China SCUM090009 KF800923 KF800933
9 Rhabdophis Jiulong, Sichuan, China GP1065 KF800924 KF800934
pentasupralabialis
10 Rhabdophis nuchalis Shennongjia, Hubei, China SICAU090001 KF800925 KF800935
11 Rhabdophis nigrocinctus Shan State, Myanmar CAS215280 KF800926 KF800936
12 Rhabdophis tigrinus Huangshan, Anhui, China GP1195 KF765390 KF765395
13 Rhabdophis tigrinus Tianshui, Gansu, China SCUM090004 KF765391 KF765396
14 Rhabdophis tigrinus Longnan, Gansu, China SCUM090001 KF765392 KF765397
15 Rhabdophis subminiatus Baoshan, Yunnan, China RDQ200905366 KF765393 KF765398
16 Rhabdophis subminiatus Panzhihua, Sichuan, China SCUM090014 KF765394 KF800927
17 Rhabdophis subminiatus Hong Kong, China SICAU090029 KF800918 KF800928

DNA extraction, amplification and sequencing. DNA was extracted from muscle tissue using a standard
phenol/chloroform extraction protocol (Sambrook et al. 1989). The polymerase chain reaction (PCR) was used for
amplification of both the cyt b and c-mos genes. For amplification of the cyt b gene, the forward primer was
L14910 (5′-GAC CTG TGA TMT GAA AAA CCA YCG TTG T-3′) according to de Queiroz et al. (2002), and the
reverse primer was H16064 (5′-CTT TGG TTT ACA AGA ACA ATG CTT TA-3′) according to Burbrink et al.
(2000). For amplification of the c-mos gene segment, the primers S77 (5′-CAT GGA CTG GGA TCAGTT ATG-3′)
and S78 (5′-CCT TGGGTG TGA TTT TCT CAC CT-3′) (Slowinski & Lawson 2002) were used. Reactions were
performed in a 25-μl reaction volume with the following cycling conditions: an initial denaturing step at 95°C for 5
min, followed by 35 cycles of 30 s at 94°C, 30 s annealing at 55–68°C, and a 1 min extension at 72°C, with a final
extension step at 72°C for 10 min. PCR was performed in the PTC-100 thermal cycler (BioRad, USA). PCR
products were purified using the DNA Agarose Gel Extraction Kit (Omega, USA) according to the manufacturer’s
instructions. Purified PCR products were sequenced on an ABI 3730 Genetic Analyzer (Applied Biosystems,
Foster City, CA, USA), using the same PCR primers. All sequences were deposited in GenBank (for accession
numbers see Table 1).
Sequence alignment and phylogenetic analysis. A total of 30 sequences were obtained for the cyt b and c-

470 · Zootaxa 3765 (5) © 2014 Magnolia Press ZHU ET AL.

mos genes; among them, Natrix natrix (Linnaeus, 1758) and Amphiesma stolatum (Linnaeus, 1758) were used as
outgroups (Table 1). The newly determined c-mos and cyt b sequences of Rhabdophis (homologous protein-coding
genes) were aligned with other retrieved sequences for the same gene regions, respectively, and to the combined
gene sequences. These three datasets were aligned using ClustalX 1.83 (Thompson et al. 1997), and translated into
amino acid sequences to check for the presence of stop codons within the sequences which might indicate the
amplification of pseudogene(s). Phylogenetic analyses derived from the combined gene fragment (c-mos + cyt b)
were performed using two different methods including Bayesian Inference (BI) and Maximum parsimony (MP).
Bayesian inference (BI) was used to reconstruct a phylogenetic bifurcating trees in the present study based on the
c-mos sequence data set. MP analyses were performed with PAUP* v4b10 (Swofford, 2003), using a heuristic
search with 1000 random stepwise additions followed by TBR branch swapping. Bootstrap branch support (BBP)
values were calculated with 1000 replicates. The best partition model was chosen using Bayes factors (BF) as
described in Brown and Lemmon (2007). BI was performed using MrBayes 3.1.2 (Huelsenbeck and Ronquist,
2001; Ronquist and Huelsenbeck, 2003), for three independent runs, each initiated with random trees. In this
analysis, posterior distributions were obtained by Markov chain Monte Carlo (MCMC) analysis with one cold
chain and three heated chains for 20 million generations and sampled every 2000th generations. The first 25% of
the trees were discarded (burn-in). Substitution parameters were unlinked and rates were allowed to vary across
partitions.
Morphological characters. One specimen was collected from Southern China and deposited at The Museum
of Biology, Sun Yat-sen University (SYS), Guangzhou, China.
Body and tail lengths were measured with a ruler to the closest 1 mm: total length (TL) from the tip of snout to
the tip of tail; snout-vent length (SVL) from the tip of snout to anterior margin of cloaca; tail length (TaL) from
posterior margin of cloaca to the tip of tail. Other measurements were taken with a caliper to the nearest 0.1 mm:
head length (HL) from the snout tip to the posterior margin of the mandible; head width (HW) at the maximal
widest part of the head; the eye horizontal diameter (ED); snout length (SL) from the tip of snout to the anterior
margin of eye; distance from eye to nostril (EN) from the anterior corner of eye to the posterior margin of nostril.
All data were collected by the same observer to avoid inter-observer bias.
The following comparative scale and maxillary teeth counts were taken: number of preoculars (PRO); number
of postoculars (PTO); number of dorsal scale rows (DSR); number of ventrals (VEN); number of subcaudals (SC);
number of maxillary teeth (MT).
Collection abbreviations: BM: The British Museum, New Oxford, London, United Kingdom; CAS: California
Academy of Sciences, San Francisco, USA; CIB: Chengdu Institute of Biology, Chinese Academy of Sciences,
Chengdu, China; DLNM: Dalian Natural History Museum, Dalian, China; GP: Peng Guo, Field tag, Sichuan,
China; HNMN: Muséum National d’Histoire Naturelle, Paris, France; IOZ: Institute of Zoology, Chinese
Academy of Sciences, Beijing, China; JK: Ke Jiang, Field tag, Sichuan, China; KIZ: Kunming Institute of
Zoology, Chinese Academy of Sciences, Kunming, China; KUZR: the Reptile collection of the Department of
Zoology, Kyoto University, Japan; LSUMZ: Louisiana State University Museum of Natural Science, Baton Rouge
City, Louisiana, USA; NXU: Ningxia University, Yinchuan, China; RDQ: Ding-qi Rao, Field tag, Kunming,
China; SCNU: South China Normal University, Guangzhou, China; SCUM: Sichuan University Museum,
Chengdu, China; SICAU: Sichuan Agricultural University, Ya’an, China; SM L: from Assistant Professor Si-Min
Lin, Department of Life Science, National Taiwan Normal University; SYS: Museum of Biology, Sun Yat-sen
University, Guangzhou, Guangdong, China; USNM: The National Museum of Natural History, Washington DC,
USA; WJC: Ji-Chao Wang, Field tag, Hainan, China; YBU: Yibin University, Sichuan, China; ZJU: Zhejiang
University, Hangzhou, China.
Specimens examined are listed in Appendix 1.

Results

Molecular phylogenetic analyses. The c-mos gene segment consisted of 588 sites, including 54 variable sites, 20
parsimony-informative (PI) sites, and 534 invariant sites. The alignment of the cyt b gene segment consisted of
1095 sites, including 436 variable sites, 325 PI sites, and 659 invariant sites. No deletions, insertions or stop codons
in the aligned sequences were detected, indicating that unintentional amplification of pseudogenes was unlikely.

A NEW SPECIES OF RHABDOPHIS FROM CHINA Zootaxa 3765 (5) © 2014 Magnolia Press · 471
New sequences generated here were deposited in GenBank (Table 1). The best-fit models for the c-mos gene
position 1, position 2, position 3, and each partition of the cyt b gene were SYM, HKY + I, HKY, GTR + I + G,
HKY + I, and GTR + G, respectively. The 50% majority consensus tree from the BI analysis of the nuDNA
fragment (c-mos) is shown in Figure 1A. Since the analyses based on the combined data yielded almost the same
topology, we show only the BI tree, but both Bayesian posterior probabilities and bootstrap values from MP are
indicated for all nodes to show the degree of congruence among results (Figure 1B).
Taxonomy. Probably due to different evolutionary characteristics of the nuclear oocyte maturation factor (c-
mos) and the mitochondrial cytochrome b (cyt b) genes, we acquired two inconsistent phylogenetic trees. But
irrespective of the topologies, our results clearly indicated that the individual from the Guangdong Province was
placed in the genus Rhabdophis with a high support (Figure 1). Fortunately, our phylogenetic analyses based on the
nuDNA (c-mos sequence data) and the combined data (c-mos + cyt b) have clearly supported the validity of this
species. The molecular results of the combined data showed that the individual shares a common ancestor with R.
adleri, R. leonardi, R. nuchalis, and R. pentasupralabialis. They form a strongly supported clade (1.00 and 80
support in Bayesian posterior probability and MP bootstrap values, respectively) that is sister to a strongly
supported clade (1.00 and 94 support in Bayesian posterior probability and MP bootstrap values, respectively)
consisting of R. swinhonis. Nevertheless, phylogenetic relationships and the evolution of the genus Rhabdophis
will have to be studied further. Combined with further evidence from morphology, the individual from Southern
China represents an undescribed species of the genus Rhabdophis. Here, we described it as a new species,
Rhabdophis guangdongensis sp. nov..

FIGURE 1. A: Bayesian phylogenetic inference tree analysis based on the c-mos sequence data set. The values on the branches
indicate posterior probability support. B: Bayesian inference tree derived from the combined gene fragment (c-mos + cyt b).
Bayesian posterior probabilities (≥90% retained) and maximum parsimony (1000 replicates) (≥50 retained) are showed at the
nodes, respectively. Numbers in brackets correspond with the ID in Table 1.

472 · Zootaxa 3765 (5) © 2014 Magnolia Press ZHU ET AL.

Rhabdophis guangdongensis sp. nov.

Holotype. Adult female, SYS r000018, collected by Ying-Yong Wang from Aizhai Village (24°56′16.58″ N,
113°39′57.82″ E; 132 m a.s.l., datum = WGS84), Renhua County, Guangdong Province, China, on May 26, 2008
(Figure 2, Figure 3: A and B).

FIGURE 2. Dorsal (A) and ventral (B) view of the holotype SYS r000018 of Rhabdophis guangdongensis sp. nov. in
preservative. Photo by Ying-Yong Wang.

FIGURE 3. The holotype SYS r000018 of Rhabdophis guangdongensis sp. nov. A: Dorsal view of head and neck in life. B:
Lateral view of head and neck in life (photo by Ying-Yong Wang); C: Dorsolateral view (Photo by Shi-Shi Lin, on Mount
Diefushan, taken on August 15, 2011).

A NEW SPECIES OF RHABDOPHIS FROM CHINA Zootaxa 3765 (5) © 2014 Magnolia Press · 473
 Diagnosis. Rhabdophis guangdongensis sp. nov. can be diagnosed by the combination of the following
characters: 1) head distinct from the neck; 2) the presence of a nuchal groove, with enlarged and paired scales on
each side; 3) maxillary teeth 20, the posterior-most three teeth abruptly enlarged, not grooved, and not separated by
diastemata from the rest of the series; 4) nostril large and lateral, in a completely divided nasal; 5) supralabials six,
the third and fourth touching the eye; 6) seven infralabials, the first four contact with anterior chin-shields; 7)
dorsal scales in 15 rows throughout, feebly keeled, the outer row smooth; 8) ventrals 126; 9) anal divided; 10)
subcaudals 39, paired; (11) the presence of a final spine at tip of tail; 12) top of head, occiput and nape before black
collar grey; 13) a broad black collar, posteriorly bordering a reverse V-shaped orange mark on the nape; 14) dorsal
surface of body and tail brown-grey, with narrow black cross-bars; 15) body and tail with two dorsolateral
longitudinal brownish-red lines, respectively with a series of white spots in cross-bars.; 16) lips with two
conspicuous black oblique bands; 17) ventral surface of head and neck cream with black spots, posteriorly
gradually becoming entirely black.
Comparisons. Comparative data of R. guangdongensis sp. nov. with 20 known species of the genus
Rhabdophis were obtained from the literature (Günther 1858, 1864; Boulenger 1893, 1900; Bourret 1935; Smith
1943; Tweedie 1957; Leviton 1970; Zhao & Jiang 1981; Zhao & Adler 1993; Zhao 1997; Zhao, et al. 1998;
Stuebing & Lian 2002; Lang & Vogel 2006; Das 2010, David & Vogel 2010) and are shown in Table 2. Examined
specimens of R. adleri, R. callichroma, R. chrysargoides, R. himalayanus, R. leonardi, R. lineatus, R. nigrocinctus,
R. nuchalis, R. pentasupralabialis, R. spilogaster, R. subminiatus, R. swinhonis, and R. tigrinus are listed in
Appendix 1.
R. guangdongensis sp. nov., R. swinhonis, R. angeli (Bourret, 1934), R. pentasupralabialis, and R. nuchalis
have the minimal number of dorsal scale rows (DSR) in this genus, equal to 15 rows throughout. The new species
is most similar to R. swinhonis, from which it differs by having 39 subcaudals vs. 44–74 in R. swinhonis; the
posterior-most three maxillary teeth strongly enlarged, not separated by diastemata from the rest of the series vs.
posterior-most two or three maxillary teeth strongly enlarged, separated by diastemata in R. swinhonis; top of head,
occiput and nape before black collar grey vs. top of head brown, occiput and nape before black collar reddish,
nuchal groove usually black and forming a longitudinal continuous or intermittent black line before black collar in
R. swinhonis; orange V-shape zone behind black collar vs. reddish zone (i.e., occiput and nape reddish) before
black collar in R. swinhonis; ventral surface of head and neck cream with black spots, posteriorly gradually
becoming entirely black vs. yellowish, powdered with brown on the sides in R. swinhonis; having dorsolateral
black cross-bars on body and tail vs. absent in R. swinhonis; and having two dorsolateral longitudinal brownish-red
parallel lines with white spots in cross-bars vs. absent in R. swinhonis. The new species differs from R. angeli by
having 20 maxillary teeth, the posterior-most three maxillary teeth strongly enlarged, not separated by diastemata
from the rest of the series vs. 22 to 23 maxillary teeth, the posterior-most two strongly enlarged, separated by
diastemata in R. angeli; top of head grey vs. brown in R. angeli; dorsal surface of body and tail brown-grey vs.
brownish in R. angeli; body and tail with two dorsolateral longitudinal brownish-red parallel lines vs. absent in R.
angeli; the presence of two series of white spots vs. a dorsolateral series of small reddish spots in R. angeli; lips
light grey vs. light brown in R. angeli; ventral surface of head and neck cream with black spots, posteriorly
gradually becoming entirely black vs. anteriorly pale orange, speckled with brown, hinder parts entirely brown in
R. angeli. The new species differs from R. pentasupralabialis by having 20 maxillary teeth, the posterior-most
three maxillary teeth strongly enlarged, not separated by diastemata from the rest of the series vs. 18 maxillary
teeth, the posterior-most two strongly enlarged, separated by diastemata in R. pentasupralabialis; top of head grey
vs. olive in R. pentasupralabialis; a broad black collar posteriorly bordering a reverse V -shaped orange zone on
the nape vs. absent in R. pentasupralabialis; lips with two conspicuous black oblique bands vs. absent in R.
pentasupralabialis; ventral surface of head and neck cream with black spots, posteriorly gradually becoming
entirely black vs. grey-white or grey-green in R. pentasupralabialis; body and tail with two dorsolateral
longitudinal brownish-red parallel lines vs. absent in R. pentasupralabialis. The new species differs from R.
nuchalis by having the posterior-most three maxillary teeth strongly enlarged, not separated by diastemata from the
rest of the series vs. the posterior-most two maxillary teeth strongly enlarged, separated by diastemata in R.
nuchalis; top of head grey vs. olive in R. nuchalis; a broad black collar posteriorly bordering a reverse V -shaped
orange zone on the nape vs. absent in R. nuchalis; lips with two conspicuous black oblique bands vs. absent in R.
nuchalis; ventral surface of head and neck cream with black spots, posteriorly gradually becoming entirely black
vs. uniform black, or black in the middle and olive on the sides in R. nuchalis; body and tail with two dorsolateral

474 · Zootaxa 3765 (5) © 2014 Magnolia Press ZHU ET AL.

longitudinal brownish-red parallel lines vs. absent in R. nuchalis.
From the remaining 16 members of the genus Rhabdophis, the new species can be readily distinguished by its
dorsal scales in 15 rows throughout, ventrals (VEN) 126, subcaudals (SC) 39 vs. 19 DSR at midbody, VEN 142,
SC 66 in R. lineatus (Peters, 1861); DSR 18(17)–17–15, VEN 149–159, SC 43–62 in R. leonardi; DSR 17–17–15,
VEN 143–162, 71–93 SC in R. auriculata (Günther, 1858); DSR 19–19–17, VEN 150–164, SC 73–88 in R. adleri;
DSR 19–19–17, VEN 165–171, SC 82–88 in R. himalayanus; DSR 19–19–17, VEN 150–170, SC 80–97 in R.
nigrocinctus; DSR 19–19–17, VEN 144–184, SC 56–97 in R. subminiatus; DSR 19–19–17 (15), VEN 144–188,
SC 38–74 in R. tigrinus; DSR 19–19–15 (17), VEN 176–185, SC 63–97 in R. murudensis (Smith, 1925); 19 DSR
at midbody, VEN 148–156, SC 75–92 in R. barbouri (Taylor, 1922); 19 DSR at midbody, VEN 152–159, SC 79–
86 in R. callichroma (Bourret, 1934); 19 DSR at midbody, VEN 138–147, SC 40–53 in R. conspicillatus (Günther,
1872); 19 DSR at midbody, VEN 143–175, SC 60–93 in R. chrysargos (Schlegel, 1837); 19 DSR at midbody, VEN
148–156, SC 75–92 in R. spilogaster (Boie, 1827); 21 DSR at midbody, VEN 156, SC 76 in R. callistus (Günther,
1873); 21 DSR at midbody, VEN 154–161, SC 64–79 in R. chrysargoides (Günther, 1858); furthermore, by outer
dorsal scale row smooth vs. keeled in R. adleri, R. barbouri, R. auriculata, R. callichroma, R. callistus, R.
chrysargoides, R. chrysargos, R. himalayanus, R. lineatus, R. murudensis, R. spilogaster; by having maxillary teeth
20 vs. more than 25 in R. auriculata, R. himalayanus, R. nigrocinctus, R. callichroma, R. chrysargos, and R. adleri.

TABLE 2. Significant characters of R. guangdongensis sp. nov. and the other known 20 species of the genus Rhabdophis
See Material and methods section for the data sources.
Species DSR Outer dorsal scale VEN SC PRO PTO MT*
row
R. guangdongensis 15–15–15 smooth 126 39 1 2 20
sp. nov.
R. angeli 16(15)–15–15 smooth 117–126 39–46 1 3 22–23
R. swinhonis 15–15–15 weakly keeled or 124–165 44–74 1 2–3 19–23(17–
smooth 21+2–3)*
R. nuchalis 15–15–15 smooth 144–169 35–65 1 3 18–22
R. pentasupralabialis 15–15–15 smooth 135–162 43–64 1 2–3 18
R. lineatus 19 at midbody strongly keeled 132–142 66–71 2 3 18(16+2)
R. leonardi 18(17)–17–15 smooth 149–159 43–62 1 2–3 19(17+2)
R. auriculata 17–17–15 strongly keeled 143–162 71–93 1–2 3 27–32
R. adleri 19–19–17 feebly keeled 150–164 73–88 1–2 3–4 27(25+2)
R. himalayanus 19–19–17 feebly keeled 165–171 82–88 2 3 26(24+2)
R. nigrocinctus 19–19–17 keeled or smooth 150–170 80–97 1 3–4 28(26+2)
R. subminiatus 19–19–17 keeled or smooth 144–184 56–97 1 3–4 23-26(21–24+2)
R. tigrinus 19–19–17(15) keeled or smooth 144–188 38–74 1–2 2–4 22-23(20–21+2)
R. barbouri 19 at midbody strongly keeled 148–156 75–92 2 3 no data
R. callichroma 19 at midbody keeled 152–159 79–86 1–2 3 27–35
R. murudensis 19–19–15(17) feebly keeled 176–185 63–97 1 3 23(21+2)
R. conspicillatus 19 at midbody keeled or smooth 138–147 40–53 1 3 no data
R. chrysargos 19 at midbody strongly keeled 143–175 60–93 1–2 3 27–35
R. spilogaster 19 at midbody feebly keeled 148–156 75–92 1–2 3–4 15
R. callistus 21 at midbody strongly keeled 156 76 1 4 no data
R. chrysargoides 21 at midbody strongly keeled 154–161 64–79 1 3 24(22+2)

*: total number of maxillary teeth (the number of maxillary teeth before the diastemata + the number of maxillary teeth
behind the diastemata).

A NEW SPECIES OF RHABDOPHIS FROM CHINA Zootaxa 3765 (5) © 2014 Magnolia Press · 475
FIGURE 4. Map showing the localities of Rhabdophis guangdongensis sp. nov. in Guangdong Province, southern China: (1)
type locality, Aizhai Village, Renhua County, Shaoguan City. (2) Mount Diefushan, Shenzhen City. (3) Mount Wutong,
Shenzhen City. (Map from http://ditu.google.cn/maps?hl=zh-CN&tab=wl, January 10, 2013)

R. guangdongensis sp. nov. is sympatric with R. subminiatus, it further differs from the latter by having two
dorsolateral longitudinal brownish-red parallel lines, respectively with a series of white spots vs. absent in R.
subminiatus; by having a broad black collar, posteriorly bordering a V-shaped orange zone on the nape vs. neck
tinged with vermilion, without broad black collar in adult R. subminiatus; the presence of a jet-black cross-bar or
triangular mark, posteriorly bordering yellow mark on the nape in young R. subminiatus.
Description of holotype. Body elongated, cylindrical; head moderately long, 3.8% of total length; tail
moderate, 16.4% of the total length; head length (HL) 1.73 times head width (HW), and significantly broader than
the neck; snout moderately long (29.7% of HL); rostral much broader than deep, just visible from above;
internasals slightly shorter than prefrontals; prefrontals wide, bending to the loreal region; frontal slightly longer
than broad, as long as the distance from the rostral to frontal; parietals longer than the frontal, barely twice as broad
in front as behind; nostril large and lateral, in a completely divided nasal; loreal single, longer than deep; eye large
(17.8% of HL), pupil rounded; supraocular single; preocular single; two postoculars; subocular absent; six
supralabials, the third and fourth touching the eye, the fifth largest; seven infralabials, the first one in contact with
each other behind the mental, first four infralabials in contact with the anterior chin-shields, the seventh
significantly extended; anterior temporal single; posterior temporal single; 20 maxillary teeth gradually posteriorly
enlarged in a continuous series, the posterior-most three teeth abruptly strongly enlarged, not grooved, and not
separated by diastemata from the rest of the series; the presence of a nuchal groove, eleven scales on each side of
groove more or less distinctly enlarged and paired; dorsal scales in 15 rows throughout, feebly keeled, the outer
most ones smooth; ventrals 126; anal divided; subcaudals 39, paired; the presence of final spine at tip of tail.
Measurements of holotype (in mm). TL 537; SVL 449; TaL 88; HL 20.2; HW 11.7; SL 6.0; ED 3.6; EN 3.2.

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 Coloration of holotype in life. Brownish-grey above; top of head, occiput and nape grey; a broad black collar
in width of five scale rows, posteriorly bordering a reverse V-shaped orange zone in width of four scale rows on the
nape; 44 pairs of narrow dorsolateral black cross-bars on body and 15 pairs on tail; body and tail with a
longitudinal brownish-red line parallel to contralateral line, respectively with a series of white spots in cross-bars;
lips light-grey with two conspicuous black oblique bands, one below the eye, another behind the eye between the
fifth and sixth supralabials; ventral surface of head and neck cream with black spots, posteriorly gradually
becoming entirely black (Figure 3: A and B).
Coloration of holotype in preservative. Grey-brown fades to gray; orange fades to yellowish-white (Figure
2)
Distribution, habitat and behaviour. Currently, R. guangdongensis sp. nov. is only known from the type
locality, Aizhai Village, Renhua County, Guangdong Province, China. On August 15, 2011, my colleague
encountered another individual of this species at Mount Diefushan (22°36′26.22″ N, 114°24′50.14″ E, 56 m a.s.l.,
datum = WGS84), Shenzhen City, Guangdong Province, China, but no sample and measurements were taken after
it was photographed (Figure 3: C). In addition, in the last three years, two individuals of this species were
photographed by tourists in submontane forest in Mount Wutong, Shenzhen City, Guangdong Province, China; one
of which was swallowing an earthworm.
Therefore, we tentatively consider that the new species is a terrestrial species distributed from the Renhua
County, northern Guangdong Province to the coastal city of Shenzhen, southern Guangdong Province, China
(Figure 4). It preys on earthworms and inhabits lowland evergreen forests and submontane forest at 56–138 m
above sea level (Figure 5).
Etymology. The specific epithet “guangdongensis” is in reference to the known distributed localities,
Guangdong Province, China. We propose the vernacular English name “Guangdong Keelback”.

FIGURE 5. Habitat of Rhabdophis guangdongensis sp. nov. at the type locality, Aizhai Village, Renhua County, Shaoguan
City, Guangdong Province. Photo by Ying-Yong Wang taken on May 26, 2008.

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 Remark. The species is considered to be locally rare. Since May 2008 to date, only the holotype of R.
guangdongensis sp. nov. was found during our extensive field surveys in southern China, including the areas from
the type locality to Shenzhen City. Only three individuals of this species were found by others. The lowland and
submontane forests are threatened by deforestation, changing land use, alien invasive plants, and ecological
degradation. Therefore, the species should be classified as a rare and endangered species in the relevant legal
provisions.

Acknowledgments

We would like to thank Peng Guo, Ding-qi Rao, Song Huang, Ke Jiang, Ji-Chao Wang and Xiao-he Wang who
offered tissue samples to Guang-Xiang Zhu. We thank the California Academy of Science, San Francisco for
kindly providing samples. We sincerely thank Dr. Gernot Vogel who provided us with morphological characters of
R. callichroma. We are very grateful to Assistant Professor Si-Min Lin for his warm help and for providing two
specimens of R. swinhonis. We also express our thanks to Yang Liu who helped us check the specimens in USA.
We extend our thanks to Xiao-he Wang, Hong-tao Cao, Zan Guo and Guo-bin Zhou et al. for their help with
fieldwork and Ping-jing Yu for help with the literature survey. We acknowledge Shi-Shi Lin for his support and for
providing us with photographs of the new species. In addition, we are much indebted to the following Institutes,
Museums and their staff for their help and permission to examine preserved specimens under their care: BM, CIB,
DLNM, HNMN, IOZ, KIZ, KUZR, NXU, SCNU, SCUM, SICAU, SYS, USNM, YBU, and ZJU.

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APPENDIX 1. Specimens examined.

Rhabdophis adleri (6): CIB10494–10495 and CIB95495, Mount Diaoluoshan, Lingshui County, Hainan Province, China;
CIB10496–10497 and CIB78032, Mount Wuzhishan, Qiongzhong County, Hainan Province, China.
Rhabdophis callichroma (2): BM1946.1.8.98(99.11.30.4), Mount Wuzhishan, Qiongzhong County, Hainan Province, China;
MNHN 1938-122 (Holotype), Mount Bavi, Tonkin.
Rhabdophis chrysargoides (3): USNM200488, Cibodas, Java, Indonesia, January 1971; USNM200489, Jakarta, Java,
Indonesia, January 1971; USNM229498, Iwahig, Palawan, Palawan Island, Philippines, February 1981.
Rhabdophis guangdongensis sp. nov. (1): SYS r000018, Aizhai Village, Renhua County, Guangdong Province, China.
Rhabdophis himalayanus (2): CIB10498, Maniweng, Medog County, Xizang Autonomous Region, China; CIB10499, Xigong
lake, Medog County, Xizang Autonomous Region, China.
Rhabdophis leonardi (6): CIB10500-04, Luding County, Sichuan Province, China, 1980; CIB14343, Pianma Town, Lushui
County, Yunnan, China.
Rhabdophis lineatus (2): USNM229280, Abachanan Barrio, Bohol Island, Philippines, April 1981; USNM318410, Leyte,
Biliran Island, Philippines, April 1987.

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 Rhabdophis nigrocinctus (3): CIB10520 and CIB10521, Xishuangbanan, Yunnan Province, China; KIZ0091, Yunnan
Province, China.
Rhabdophis nuchalis (14): CIB10524–25, CIB10528, CIB10539, CIB10552, CIB10564 and CIB10588, Lichuan City, Hubei
Province, China; SICAU090005-090007 and SICAU090014, Tianquan County, Ya’an City, Sichuan Province, China,
August 2009; SCUM090005 and SCUM090007–090008, Xingou Town, Ya’an City, Sichuan Province, China, August
2009.
Rhabdophis pentasupralabialis (7): CIB10865, Mount Emei, Sichuan Province, China; CIB10713, CIB10714, CIB10812,
CIB10818 and CIB10853, Jiulong County, Sichuan, China; JK090501, Ke Jiang Field tag, Jiulong County, Sichuan,
China, May 2009.
Rhabdophis spilogaster (2): USNM498930, Baggao Municipality, Cagayan Province, Luzon Island, Philippines, May 1989;
USNM499859, Tagaytay Ridge, Batangas Province, Luzon Island, Philippines, April 1990.
Rhabdophis subminiatus (29): SYS r000083, Heishiding National Nature Reserve, Guangdong Province, China, September
2009; SYS r000085, Renhua County, Guangdong Province, China, September 2009; SYS r000409, Shimentai Nature
Reserve, Yingde City, Guangdong Province, China, September 9, 2011; SYS r000479–0480, Mount Yinpingshan,
Dongguan City, Guangdong Province, China, December 18, 2011; SYS r000751, Renhua County, Guangdong Province,
China, September 2012; ZJU Cm1, Fujian Province, China, October 1963; SCUM090013–14, Miyi County, Panzhihua
City, Sichuan Province, China, August 2009; IOZ74007, Mojiang County, Yunnan Province, China; CIB10978,
IOZ75018, IOZ79003, IOZ79019 and IOZ79058, Pingbian County, Yunnan Province, China. KIZ69 I 0007 and KIZ69
I 1004, Jinggu County, Yunnan Province, China; KIZ69 I 0026, Shuangjiang County, Yunnan Province, China;
CIB10916-10918, Luodian County, Guizhou Province, China; CIB10877-10879, CIB10884, CIB10889 and CIB78047,
Diaoluoshan, Lingshui County, Hainan Province, China; CIB10885 and CIB10887, Qiongzhong County, Hainan
Province, China.
Rhabdophis swinhonis (5): SCUM090015, Mount Dahan, Pingtung County, Taiwan, spring 2008; KUZR 18977, Taiwan;
CIB72588, Xiaogui Lake, Huotai Town, Pingtung County, Taiwan; SM L001 and SM L002, Taiwan, 2013.
Rhabdophis tigrinus (29): SYS r000062, Mount Sanqing, Jiangxi Province, China, June 2007; SYS r000025 and 0033,
Yangjifeng Nature Reserve, Jiangxi Province, China, June 2008; CIB11056–57, Huoshan County, Anhui Province, China;
CIB11279, Youyang County, Chongqing City, China; CIB11280, Gulin County, Sichuan Province, China; CIB78545–46,
Panshi City, Jilin Province, China; SCUM090001, Huixian County, Gansu Province, China, July 2009; SCUM090003–4,
Tianshui County, Gansu Province, China, July 2009; SICAU040001, Mount Liupanshan, Ningxia Province, China,
August 2004; SCUM090024, Taiwan; SICAU050001, Mount Liupanshan, Ningxia Province, China, August 2005;
KIZ89 II 031–32, Youyang County, Chongqing City, China, 1989; IOZH10735, Miaoershan Nature Reserve, Guilin City,
Guangxi Province, China; IOZ00408, Baiyangdian, Baoding City, Hebei Province, China; ZJU03, ZheJiang Province,
China; DLNM632296 and DLNM632400, Dandong City, Liaoning Province, China; NXU09, Mount Helanshan, Ningxia
Province, China; GP1195–1200, Peng Guo Field tag, Huangshan City, Anhui Province, China, July 2009.

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