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Revisions of two poorly known species of Opisthotropis Günther, 1872 (Squamata: Colubridae: Natricinae) with description of a new species from China

March 2017
Zootaxa 4247(4):391-412

DOI:10.11646/zootaxa.4247.4.3

Authors:

Ying-Yong Wang

Sun Yat-sen University

Zuyao Liu

University of Bern

Zhi-Tong Lyu

Chinese Academy of Sciences

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The previous descriptions of Opisthotropis maxwelli Boulenger, 1914 and O. andersonii (Boulenger, 1888) were considered imperfect due to the limited number of specimens. This may in turn cause a problem for accurate species identification. In our study, the species boundaries of these two species were investigated using an integrative approach incorporating morphological characters and molecular phylogenetic analyses of the mitochondrial Cyt b gene of 26 specimens of nine known Opisthotropis species collected from southeastern China. Our results surprisingly revealed a new cryptic species, Opisthotropis shenzhenensis sp. nov., from Shenzhen and Dongguan, Guangdong Province, southern China. Further, we re-described O. maxwelli based on several specimens from Fujian and Guangdong, and O. andersonii based on a series of specimens from Hong Kong, Shenzhen and Guangzhou, China. We provide an updated identification key to all described Opisthotropis species from China.

. Specimens used for molecular analyses in this study.

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Bayesian inference tree of relationships among mitochondrial Cyt b nucleotide seqeucnes from Opisthotropis. Numbers above branches (>70% retained) indicate Bayesian posterior probabilities and numbers below branches (>60 retained) are bootstrap support for maximum likelihood (500 replicates) analyses.

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General aspect of Opisthotropis andersonii. A: dorsolateral view of the adult female specimen SYS r001020 in life. B: Ventral view of the specimen SYS r001020. C and D: Adult male specimen SYS r000893 in life. E: Lateral view of adult female SYS r001382. Photos by JW, JZ, JHY and ZYL.

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General aspect and scalation of head of Opisthotropis shenzhenensis sp. nov.. A: Dorsal view of the holotype SYS r001018. B: Ventral view of the holotype. C: Dorsolateral view of head and anterior neck of the holotype in life. D: General aspect of SYS r001145 in life. Photos by JW and JZ. Opisthotropis shenzhenensis sp. nov., O. andersonii, O. lateralis and O. kuatunensis are sympatric species from Mt. Wutong, Shenzhen. O. shenzhenensis differs from O. lateralis by having DSR 19:19:19 vs. 17:17:17; a single preocular vs. two preoculars; uniformly olive-green above with black mesh pattern on dorsal surface of body and tail vs. brown above, with a distinct longitudinal black stripe on both sides of body. O. shenzhenensis differs from O. kuatunensis by having 9–10 supralabials, all of which entire vs. 13–16 supralabials, first six to eight entire, posterior ones divided horizontally; dorsal scales weakly keeled on neck, strongly keeled on body and tail vs. extremely strongly keeled throughout; olive-green above with black mesh pattern on dorsal body and tail vs. brown above, with black longitudinal stripes on body and tail. Opisthotropis shenzhenensis sp. nov. can be easily distinguished from other Chinese congeners as follows: It differs from O. latouchii in having DSR 19:19:19 vs. 17:17:17; loreal not entering the orbit vs. entering the orbit; olive-green above with black mesh pattern vs. yellowish green above, with alternate longitudinal black and yellow stripes. It differs from O. cheni in having DSR 19:19:19 vs. 17:17:17; loreal not entering the orbit vs. entering the orbit; olive-green above with black mesh pattern vs. dark olive brown above, with yellow crossbars or marks. It differs from O. laui in having DSR 19:19:19 vs. 25:23:23; olive-green above with black mesh pattern vs. dark olive above, with pale yellow crossbars. It differs from O. balteata in having DSR 19:19:19 vs. 19:19:17; V 162–179, SC

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Comparisions of head scalation of a: Opisthotropis shenzhenensis sp. nov. (holotype SYS r001018), b: O. maxwelli (SYS r000841), c and d: O. andersonii (SYS r001020, SYS r001082). Line illustration by ZTL.

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Accepted by T. Nguyen: 7 Feb. 2017; published: 29 Mar. 2017

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN

1175-5334

(online edition)

Zootaxa 4247 (4): 391

–

412

http://www.mapress.com/j/zt/

Article

391

https://doi.org/10.11646/zootaxa.4247.4.3

http://zoobank.org/urn:lsid:zoobank.org:pub:B926664F-8D4E-4CBA-8EEA-317B5AB46357

Revisions of two poorly known species of Opisthotropis Günther, 1872

(Squamata: Colubridae: Natricinae) with description of a new species from

China

YING-YONG WANG

1,4

, QIANG GUO

, ZU-YAO LIU

, ZHI-TONG LYU

, JIAN WANG

, LIN LUO

YAN-JUN SUN

& YAN-WU ZHANG

State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275,

P.R. China

Shenzhen Administrative Bureau for Wildlife and Forest Plants Protection, Shenzhen 518048, P.R. China

Shenzhen Park Service, Shenzhen 518040, P.R. China

Corresponding author. E-mail: wangyy@mail.sysu.edu.cn

Abstract

The previous descriptions of Opisthotropis maxwelli Boulenger, 1914 and O. andersonii (Boulenger, 1888) were consid-

ered imperfect due to the limited number of specimens. This may in turn cause a problem for accurate species identifica-

tion. In our study, the species boundaries of these two species were investigated using an integrative approach

incorporating morphological characters and molecular phylogenetic analyses of the mitochondrial Cyt b gene of 26 spec-

imens of nine known Opisthotropis species collected from south-eastern China. Our results surprisingly revealed a new

cryptic species, Opisthotropis shenzhenensis sp. nov., from Shenzhen and Dongguan, Guangdong Province, southern Chi-

na. Further, we re-described O. maxwelli based on several specimens from Fujian and Guangdong, and O. andersonii

based on a series of specimens from Hong Kong, Shenzhen and Guangzhou, China. We provide an updated identification

key to all described Opisthotropis species from China.

Key words: China, mitochondrial Cyt b, morphology, Opisthotropis andersonii, Opisthotropis maxwelli, Opisthotropis

shenzhenensis sp. nov., taxonomy

Introduction

The genus Opisthotropis Günther, 1872, which currently comprises 21 species, has a widespread distribution

throughout southern China and mainland of Southeast Asia, eastward to the Ryukyu Archipelago of Japan,

southward to Sumatra in Indonesia and the Philippines (Ziegler et al. 2008; David et al. 2011, 2015; Yang et al.

2013; Teynié et al. 2014). Opisthotropis are mostly nocturnal, and inhabit flowing streams and occasionally dwell

in riparian habitats (Yang et al. 2013). Many species were described from a single or a few specimens, and still are

known imperfectly (Stuart & Chuaynkern 2007; Ziegler et al. 2008; David et al. 2011; Yang et al. 2013; Teynié et

al. 2014). For example, O. maxwelli (Boulenger, 1914), a rare species endemic to southeastern China, was

described originally from a single female specimen, and re-described based on six old, faded specimens (Zhao et

al. 1998). Thus, the pattern and coloration in life are not known. Opisthotropis andersonii (Boulenger, 1888) is

known to occur in Hong Kong, Shenzhen, China and Vietnam (Zhao et al. 1998; David et al. 2011). This species

was described originally from a single juvenile specimen and was re-described based on two specimens deposited

in the Natural History Museum, London (Zheng 1992). In the past few years, during extensive herpetological

surveys conducted in southern China, we found that the morphological characters of some O. andersonii specimens

were not consistent with the diagnostic characters given in previous descriptions, for example, in half of the

specimens collected the loreal enters the orbit, as opposed to loreal not entering the orbit in the previous

descriptions (Boulenger 1888; Zheng 1992; Zhao et al. 1998). Therefore, revisions of the diagnostic characters of

O. maxwelli and O. andersonii are imperative for accurate species identification.

WANG ET AL.

392

During the field surveys conducted in mountainous areas, Shenzhen, Guangdong Province, China, we

collected four sympatric species of Opisthotropis, namely O. andersonii, O. kuatunensis Pope, 1928, O. lateralis

Boulenger, 1903, and another unnamed species, which is most similar to O. andersonii in habitus and color pattern.

The unnamed Opisthotropis can be distinguished from other congeners by distinctive morphological differences,

from the nine known congeners of China by significant molecular differences in the mitochondrial Cyt b gene.

Therefore, we herein describe it as a new species.

Material and methods

Molecular analyses. We analysed a total of 28 samples from nine of the 11 currently recognized species of

Opisthotropis from China (tissue from O. balteata (Cope, 1895) and O. jacobi Angel & Bourret, 1933 was not

available), an unnamed Opisthotropis taxon from Shenzhen and Dongguan, Guangdong Province, China; all

collected localities of samples of Opisthotropis were marked in Fig.1; Rhabdophis guangdongensis Zhu, Wang,

Takeuchi & Zhao, 2014 and Hebius craspedogaster (Boulenger, 1899) were used as the out-groups. Details of

samples sequenced for mitochondrial Cyt b and their associated GenBank accession numbers are presented in

Table 1.

DNA was extracted from liver tissue using a standard phenol-chloroform extraction protocol (Sambrook et al.

1989). The mitochondrial Cyt b gene was PCR amplified and sequenced using the primer L14919 (5’-

AACCACCGTTGTTATTCAACT-3’) and H16064 (5’-CTTTGGTTTACAAGA ACAATGCTTTA-3’) (Burbrink

et al. 2000; Guo et al. 2012). PCR amplifications were performed in a 25µL reaction volume with the following

cycling conditions: an initial denaturing step at 94°C for 7 min; 40 cycles of denaturing at 94°C for 40 s, annealing

at 46°C for 30 s and extending at 72°C for 1 min, and a final extending step of 72°C for 8 min. PCR products were

purified with spin columns. The purified products were sequenced with both forward and reverse primers using

BigDye Terminator Cycle Sequencing Kit according to the guidelines of the manufacturer. The products were

sequenced on an ABI Prism 3730 automated DNA sequencer in Shanghai Majorbio Bio-pharm Technology Co.,

Ltd.

We obtained 28 sequences of the 1100 bp of the Cyt b gene in this study, among which DNA data of six

samples were obtained from GenBank (Table 1). Alignments were first conducted using Clustal X 2.0 (Thompson

et al. 1997), with default parameters and then the alignment was checked and manually revised. The HKY model

(Hasegawa 1985), assuming a gamma shaped distribution across sites (Felsenstein 2004), was selected as the best-

fitting nucleotide substitution model using model test module in MEGA 6 (Tamura et al. 2013). Maximum

likelihood (ML) was implemented in MEGA 6 and Bayesian inference (BI) using MrBayes 3.12 (Ronquist &

Huelsenbeck 2003). The ML and BI methods were chosen to conduct phylogenetic analyses. For ML analysis, the

bootstrap consensus tree inferred from 1000 replicates was used to represent the evolutionary history of the taxa

analyzed. Branches corresponding to partitions reproduced in less than 60% of bootstrap replicates were collapsed.

For BI analysis, two independent runs with four Markov Chain Monte Carlo simulations were performed for one

million iterations and sampled every 100th iteration. The first 25% of samples were discarded as burn-in.

Convergence of the Markov Chain Monte Carlo simulations was assessed using Tracer v.1.4 (http://

tree.bio.ed.ac.uk/ software/tracer/). Apart from phylogenetic tree-based methods, we also calculated pairwise

sequence divergence based on uncorrected p-distance implemented in MEGA 6.

Morphological characters. Measurements of head, body and tail were taken with a digital caliper to the

nearest 1 mm, and of loreal and rostral to the nearest 0.1 mm: total length (TL) from the tip of the snout to the tip of

the tail; snout-vent length (SVL) from the tip of the snout to the anterior margin of the cloaca; tail length (TaL)

from the posterior margin of the cloaca to the tip of the tail; head length (HL) from the tip of the snout to the

posterior margin of the mandible; maximum head width (HW); maximum rostral width (RW); maximum loreal

depth (LoD); maximum loreal length (LoL). Scalation features and abbreviations are as follows: internasals (IN);

prefrontal (PrF); preoculars (PrO); postoculars (PtO); loreals (L); supralabials (SPL); infralabials (IFL); temporals

(TMP); ventral scales (V); subcaudals (SC); dorsal scale rows (DSR) were counted at one head length behind head,

at midbody, and at one head length before vent respectively. Bilateral scale counts were given as left/right.

Comparative data for Opisthotropis shenzhenensis sp. nov. and 21 congeners were obtained from literature

(Günther 1872; Boulenger 1893; Taylor 1965; Orlov et al. 1998; Zheng 1992; Zhao et al. 1998; Zhao 2004, 2006;

393

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

Stuart & Chuaynkern 2007; Murphy et al. 2008; Ziegler et al. 2008; Iskandar & Kamsi 2009; David et al. 2011;

and Yang et al. 2011, 2013; Nguyen et al. 2014). The voucher details for the seven species of Opisthotropis that we

examined are listed in the Appendix.

Collection abbreviations: SYS, The Museum of Biology, Sun Yat-sen University, Guangzhou, China. YBU,

Yibin University, Yibin, China. FJNU, The Biology Museum, Fujian Normal University.

TABLE 1. Specimens used for molecular analyses in this study.

No. Species name Map location code: Country: Location Voucher No. Cyt b gene

(1) O. shenzhenensis sp. nov. 2: China; Mt. Wutong, Shenzhen, Guangdong SYS r001018 KY594727

(2) O. shenzhenensis sp. nov. 3: China: Sanzhoutian, Shenzhen City, Guangdong SYS r001021 KY594728

(3) O. shenzhenensis sp. nov. 4: China: Mt. Tiantou, Shenzhen City, Guangdong SYS r001032 KY594729

(4) O. andersonii 1: China: Tai Tam, HK Island, Hong Kong SYS r001423 KY594730

(5) O. andersonii 1: China: Tai Mo Shan, Hong Kong SYS r001424 KY594731

(6) O. andersonii 2: China: Mt. Wutong, Shenzhen City, Guangdong SYS r001020 KY594732

(7) O. andersonii 2: China: Mt. Wutong, Shenzhen City, Guangdong SYS r001082 KY594733

(8) O. andersonii 6: China: Mt. Maofeng, Guangzhou City, Guangdong SYS r001382 KY594734

(9) O. andersonii 6: China: Mt. Maofeng, Guangzhou City, Guangdong SYS r001383 KY594735

(10) O. maxwelli 7: China: Nan’ao Island, Nan’ao County, Guangdong SYS r000841 KY594736

(11) O. maxwelli 8: China: Huboliao Nature Reserve, Nanjing County,

Fujian

SYS r001053 KY594737

(12) O. laui 14: China: Shangchuan Island, Taishan County,

Guangdong

SYS r001161 KY594738

(13) O. laui 14: China: Shangchuan Island, Taishan County,

Guangdong

SYS r001170 KY594739

(14) O. laui 14: China: Shangchuan Island, Taishan County,

Guangdong

SYS r001171 KY594740

(15) O. cheni China: Guangdong YBU071040 GQ281779

(16) O. cheni 12: China: Shimentai Nature Reserve, Yingde City,

Guangdong

SYS r001422 KY594741

(17) O. latouchii 10: China: Guadun Village (=Kuatun), Wuyishan City,

Fujian

SYS r000670 KY594742

(18) O. latouchii China: Fujian - GQ281783

(19) O. lateralis 13: China: Heishiding Nature Reserve, Fengkai County,

Guangdong

SYS r000951 KY594743

(20) O. lateralis 2: China: Mt. Wutong, Shenzhen, Guangdong SYS r001080 KY594744

(21) O. lateralis China: Guangxi - GQ281782

(22) O. kuatunensis 11: China: Qixiling Nature Reserve, Yongxin County,

Jiangxi

SYS r000998 KY594745

(23) O. kuatunensis 9: China: Wulong Village, Shanghang County, Fujian SYS r001008 KY594746

(24) O. kuatunensis 2: China: Mt. Wutong, Shenzhen City, Guangdong SYS r001081 KY594747

(25) O. guangxiensis China: Guangxi - GQ281776

(26) O. maculosa 13: China: Heishiding Nature Reserve, Fengkai County,

Guangdong

SYS r000946 KY594748

(27) Rhabdophis

guangdongensis

China: Aizhai Village, Renhua County, Guangdong SYS r000018 KF800930

(28) Hebius craspedogaster China: Xizang YBU071128 GQ281781

WANG ET AL.

394

FIGURE 1. Sampling localities of Opisthotropis in China. 1: Hong Kong; 2: Mt. Wutong; 3: Sanzhoutian; 4: Mt. Tiantou; 5:

Mt. Yinping; 6: Mt. Maofeng; 7: Nan’ao Island; 8: Huboliao Nature Reserve; 9: Wulong Village; 10: Guadun Village; 11:

Qixiling Nature Reserve; 12: Shimentai Nature Reserve; 13: Heishiding Nature Reserve; 14: Shangchuan Island. The red dot

represent localities of Opisthotropis shenzhenensis sp. nov.. See Table 1 for locality codes and detailed locality information.

Results

Phylogenetic analyses. The Maximum likelihood (ML) and Bayesian inference (BI) phylogenetic trees were

constructed based on concatenated DNA sequences of the mitochondrial Cyt b gene with a total length of 1100-bp.

Two analyses resulted in essentially identical topologies with strongly node supporting values, except the clade

composed of Opisthotropis guangxiensis and O. maculosa. The phylogeny tree (Fig. 2) indicated that the

populations of Opisthotropis from Shenzhen and Hong Kong are divided into two strongly supported monophyletic

clades (PP = 1.00, BP = 72) with a deep mtDNA sequence divergence (p-distance 0.109–0.117, see Table 2), which

were at a same divergence level between above two clades and O. maxwelli (p-distance 0.107–0.124), whereas the

smallest p-distance between two recognized species O. cheni and O. latouchii was 0.050. Therefore, these two

clades represent two taxa. Furthermore, the first clade is identified as O. andersonii, because specimens within this

clade, including specimens SYS r001423 and SYS r001424 from its type locality Hong Kong, and SYS r001020,

SYS r001082, SYS r001382 and SYS r001383 from Guangdong, possess the following characters: DSR 17:17:17,

a single loreal, more than twice as long as deep, dorsal body scale rows weakly keeled; another clade is composed

of specimens SYS r001021, SYS r001018 and SYS r001032, which possess DSR 19:19:19, loreal short, 1.3–1.6

times as long as deep, dorsal body scale rows rather strongly keeled, therefore, it represents an undescribed species.

Herein, we re-describe O. maxwelli and O. andersonii based on newly collected specimens, and describe a new

species, Opisthotropis shenzhenensis sp. nov.

395

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

7$%/(8QFRUUHFWHGSGLVWDQFHVDPRQJWHQVSHFLHVRI OpisthotropisDQGRXWJURXSVEDVHGRQD&\WbIUDJPHQW

1R 6SHFLHV              

 O. shenzhenensis VSQRY 

 O. shenzhenensis VSQRY 

 O. shenzhenensis VSQRY 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. andersonii             

 O. andersonii             

 O. maxwelli             

 O. maxwelli             

 O. laui             

 O. laui             

 O. laui             

 O. cheni             

 O. cheni             

 O. latouchii             

 O. latouchii             

 O. lateralis             

 O. lateralis             

 O. lateralis             

 O. kuatunensis             

 O. kuatunensis             

 O. kuatunensis             

 O. guangxiensis             

 O. maculosa             

 Rhabdophis guangdongensis             

 Hebius craspedogaster             

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WANG ET AL.

396

7$%/(&RQWLQXHG

1R 6SHFLHV               

 O. shenzhenensis VSQRY 

 O. shenzhenensis VSQRY 

 O. shenzhenensis VSQRY 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. andersonii 

 O. maxwelli 

 O. maxwelli 

 O. laui 

 O. laui 

 O. laui 

 O. cheni 

 O. cheni 

 O. latouchii 

 O. latouchii 

 O. lateralis 

 O. lateralis 

 O. lateralis 

 O. kuatunensis 

 O. kuatunensis 

 O. kuatunensis              

 O. guangxiensis              

 O. maculosa              

 Rhabdophis guangdongensis              

 Hebius craspedogaster              

397

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

FIGURE 2. Bayesian inference tree of relationships among mitochondrial Cyt b nucleotide seqeucnes from Opisthotropis.

Numbers above branches (>70% retained) indicate Bayesian posterior probabilities and numbers below branches (>60 retained)

are bootstrap support for maximum likelihood (500 replicates) analyses.

Systematics

Opisthotropis maxwelli Boulenger, 1914

Figs. 3 and 6b

Opisthotropis maxwelli Boulenger, 1914. Type locality: “South Forkien” (= Fujian)

Specimens examined. Three specimens: SYS r001273, adult female, from Wulong Village (25°12′48.33″N,

116°50′10.69″E; 995 m a.s.l.), Shanghang County, Fujian Province, collected by Jian Wang (JW hereafter), Ying-

Yong Wang and Hai-Long He on 16 July 2015; SYS r000841, subadult female, from Nan’ao Island (23°26′0.09″N,

117°4′45.61″E; 425 m a.s.l.), collected by Jian Zhao (JZ hereafter) and Yu-Long Li (YLL hereafter) on 30 June

2013; SYS r001053, adult male, from Huboliao Nature Reserve (24°34'32.26"N, 117°13'40.61"; 739 m a.s.l.) by

Zu-Yao Liu (ZYL hereafter) and Zhi-Tong Lyu (ZTL hereafter) on 22 September 2014; further information of

collection localities is listed in Table 1. Five previously collected specimens in buffered formalin from the College

of Life Sciences, Fujian Normal University: FJNU 3611001, 3611003, 3611004, 3611005 and 3611007, all

collected from Nanjing County, Fujian Province, China. The specimens FJNU 3611001, 3611003 and 3611004

were already described by Zhao et al. (1998) and numbered as 1045, 1433 and 561, respectively.

Description. Measurements, scalation and body proportions of six specimens are listed in Table 3. The

examined specimens roughly conform to the previous descriptions (Boulenger 1914; Zhao et al. 1998), and show

WANG ET AL.

398

some variation. Body cylindrical; TL 319–442 mm; tail relatively long, TaL 20–23% of TL; head barely distinct

from the neck; snout short, broad and depressed; rostral visible from above, approximately twice as broad as deep,

RW 28–32% of HW; nasal directed dorsolaterally, its lower edge inverted V-shaped, bordered by first and second

supralabials ventrally, by loreal and prefrontal posteriorly, by internasal dorsally, by rostral anteriorly; a short

vertical nasal cleft below the nostril, pointing to the apex angle of first supralabial and dividing nasal into anterior

and posterior parts; supralabials 7/7 (n=7), 7/6 (n=1), the first supralabial pentagonal, usually fifth entering the

orbit (the fourth entering the orbit in FJNU 3611004, fourth and fifth entering the orbit in right side of head in SYS

r001273), the last one (seventh or sixth) longest, longer than total length of two preceding supraliabials; loreal

single, 1.4–1.7 times as long as deep, not touching internasal, not entering the orbit, usually in contact with second,

third and fourth supralabials (with third and fourth in FJNU 3611001); two (rarely one) preoculars; two

postoculars; temporals 1+2/1+2 (n=2), 1+1/1+1 (n=5), 1+1/1+2 (n=1), anterior temporal significantly elongate;

infralabials 8/8, first four or five touching anterior chin-shields on both sides; two pairs of chin-shields, the anterior

pair large, in contact with each other medially; posterior chin-shields significantly small, separated from each other

by small scales; dorsal scale rows 17:17:17; dorsal scales smooth on the anterior neck (dorsal scales on neck

smooth in the previous descriptions (Boulenger 1914; Zhao et al. 1998)), presenting barely perceptible weak keel

on sixth vertebral scale in SYS r000841, on ninth vertebral scale in FJNU 3611005 and SYS r001273, on 11th

vertebral scale on FJNU 3611004, on 12th vertebral scale in FJNU 3611007, on 15th vertebral scale in FJNU

3611003, on 16th vertebral scale in SYS r001053, succeeding dorsal scales feebly keeled, rather strongly keeled on

posterior part of body, but most outer one or two scale rows smooth; V 149–157; cloacal plate divided; subcaudals

54–62, paired (V 147–155, SC 53–61 in descriptions by Zhao et al. (1998)).

Coloration in life. Dorsal surface of head and neck olive-green in SYS r001053 and SYS r001273, with black

blotches on each scales; dorsal surface of body and tail olive-green with intermittent longitudinal black stripes

parallel to each other, crossing each scale; the outer most dorsal scale row yellow, occasionally with black spots;

mental and infralabials black; venter yellow, chin-shields, throat and subcaudal series speckled with black. The

specimen SYS r000841 is similar to SYS r001053 in color and pattern in life, but its dorsal surface of head, body

and tail dark olive-brown, with 17 bright yellow spots on anterior part of body and barely perceptible longitudinal

black stripes, crossing each scale; lower part of the outer most dorsal scale row yellow, with small blackish grey

spots; ventral surface yellow, mottled with blackish grey on chin-shields, throat and subcaudal series.

Coloration in preservative. In ethanol, blackish-brown or blackish-grey above, longitudinal stripes of dorsal

body visible in SYS r001053, not visible in SYS r000841 and 1273; yellowish beneath. In buffered formalin,

brown above, longitudinal stripes distinct, two ventrolateral stripes especially eye-catching; whitish beneath.

Revision of diagnosis. This species is characterized by the combination of (1) TL 319–442 mm; (2) tail

relatively long, TaL 20–23% of TL; (3) rostral small, RW 28–32% of HW; (4) nasal in contact with first and second

supralabials ventrally; (5) nasal cleft invariably pointing to the apex angle of first supralabial; (6) loreal not

entering orbit, 1.4–1.7 times as long as deep; (7) supralabials 6–7, the last one longest; (8) infralabials 8; (9) dorsal

scale rows 17:17:17; (10) dorsal scales smooth on the anterior neck, those following feebly keeled, rather strongly

keeled on posterior part of body up to the tail; (11) ventrals 147–157, subcaudals 53–62; (12) olive-brown to olive-

green above in life, body and tail with distinct or barely perceptible longitudinal black stripes, crossing each scale;

yellow beneath, ventral surface of head and tail mottled with blackish grey.

Distribution and habitat. According to the known specimens and literature (Boulenger 1914; Zheng 1992,

Zhao et al. 1998; Zhong 2004), O. maxwelli is distributed in the southeast of Guangdong Province, the southwest

of Fujian Province and south of Jiangxi Province. It inhabits montane slow-flowing streams with sand and gravel

bottom at elevations between 425–1000 m a.s.l.

Remarks. Opisthotropis maxwelli was described on the basis of a single female specimen from the British

Museum (now the Natural History Museum, London), presented by J. P. Maxwell (Boulenger, 1914), and specimen

number was not given. Zhao et al. (1998) described and measured six specimens of O. maxwelli, of which,

specimens FJNU 1433 and 1045 measured at 610 mm and 520 mm in SVL, respectively. However, we obtained

measurements of 310 mm and 405 mm in SVL, respectively.

399

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

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/R//R' ± ± ± ± ± ±

WANG ET AL.

400

FIGURE 3. General aspect of Opisthotropis maxwelli. A and B: Adult male specimen SYS r001053. C: Dorsolateral view of

the subadult female specimen SYS r000841 in life. D: Adult female specimen SYS r001273 in life. Photos by JW, JZ and ZYL.

Opisthotropis andersonii (Boulenger, 1888)

Figs. 4 and 6c, d

Calamohydrus Andersonii Boulenger, 1888. Type locality: Hong Kong.

Opisthotropis andersonii: Boulenger 1893

Opisthotropis andersoni: Smith 1943

Opisthotropis andersonii: Zheng 1992; Zhao et al. 1998; Yang et al. 2013

Specimens examined. Ten specimens from China: SYS r001423, adult female, collected by Anthony Lau on 29

January, 2015, from Tai Tam (22°15′32.52″N, 114°12′6.6″E; 252 m a.s.l.); SYS r001424, adult female, collected by

Yik-Hei Sung on 20 May, 2015, from Tai Mo Shan (22°24′36.18″N, 114°7′12.12″E; 740 m a.s.l.). SYS r000607,

adult male, collected by Jian-Huan Yang (JHY hereafter) and Run-Lin Li (RLL hereafter) on 8 March, 2012; SYS

r001017, juvenile, and SYS r001020, adult female, collected by JZ and RLL on 15 and 16 September, 2014; SYS

r001082, adult male, collected on 13 March 2015 by JW and ZTL, all from Mt. Wutong (22°34′54.8″N,

114°12′2.7″E; 250–300 m a.s.l.). SYS r000803, adult male, collected by JZ and YLL on 5 April, 2013 from Mt.

Yangtai (22°38′36.09″N, 113°59′7.45″E; 150 m a.s.l.), Shenzhen City, Guangdong Province. SYS r000893, adult

male, collected by JZ and YLL on 3 November, 2013 from Dapeng Peninsula (22°30′17.55″N, 114°32′23.01″E;

102 m a.s.l.), Shenzhen City, Guangdong Province. SYS r001382, an adult female, and SYS r001383, a juvenile,

by JW on 16 October, 2015 from Mt. Maofeng (23°18′22.79″N, 113°22′58.03″E; 55 m a.s.l.). Further information

of collection localities is listed in Table 1.

Description. Measurements, scalation and body proportions of eight adult specimens are listed in Table 3. The

examined specimens roughly conform to the previous descriptions (Boulenger 1888; Zhao et al. 1998), and show

401

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

significant variation. Body cylindrical; adult specimens measuring 378–462 mm in TL; tail short, TaL of 15–20%

of TL; Head small, indistinct from neck; rostral concave, visible from above, slightly enlarged, more than twice as

broad as deep, RW 31–37% of HW; nasal directed dorsolaterally, rectangular or pentagonal, in contact with first

two supralabials ventrally, with loreal and prefrontal posteriorly, with internasal dorsally, with rostral anteriorly; a

short vertical cleft below the nostril and dividing nasal into anterior and posterior parts, pointing to the posterior

upper-angle of first supralabial (n=3), to the anterior upper-angle of second supralabial (n=3), or connecting with

the suture between first and second supralabials (n=4); a single loreal, more than twice as long as deep (LoL/LoD

2.1–2.9), not in contact with internasal, in contact with second, third and fourth supralabials (in contact with

second, third, fourth and fifth in left side of head of SYS r000803), whereas only in contact with third and fourth

supralabials in original description (Boulenger 1888); in specimens SYS r001423, SYS r001020, 0893 and 0607,

loreal not entering the orbit, preoculars 2–3 ( 3 only on right side in SYS r001423), upper one significantly larger

than the lower ones; in SYS r001424, SYS r000803, 1382, 1082, 1017 and 1383, loreal entering the orbit, preocular

absent in SYS r000803, 1382 and 1383, present a small preocular under the loreal in SYS r001424, SYS r001082

and SYS r001017; postoculars 1–2; anterior temporal single, significantly prolonged (anterior temporal broken into

three in specimen SYS r001423); posterior temporals two, only one on left side of head in specimen SYS r001423;

supralabials 8/8 in all adults, 7/7 in two juveniles, commonly first three rectangular, the last one significantly

smaller than the adjacent preceding supralabial; the fifth, or fourth and fifth, or fourth, fifth and sixth supralabials

touching the orbit; 7–10 infralabials, the first one in contact with its fellow behind the mental; two pairs of chin

shields; anterior chin-shields large, in contact with each other, and in contact with the first four or first five

infralabials on both sides; posterior chin shields small and separated by small scales; dorsal scale rows 17:17:17;

dorsal scales weakly keeled throughout the body in previous description (Zhao et al. 1998; Teynie et al. 2014),

whereas smooth on the anterior neck in our specimens, first presenting barely perceptible weak keel on 10

vertebral scale in SYS r001423, on 13

in SYS r001424, 15

in SYS r000803 and 1382, 17

in SYS r001020, 18

in SYS r001082, 21

in SYS r000607, 30

in SYS r001017, 46

in SYS r001383, 58

in SYS r000893, and

gradually visible posteriorly; dorsal scales moderately and visibly keeled at mid-body to posterior part of body, the

outermost row smooth; dorsal scales rather strongly keeled on tail; V 141–174; cloacal plate divided; subcaudals

43–59, paired.

Coloration in life. Dorsal surfaces of head, body and tail olive-green to olive-brown, usually each scale with

black lateral edges, forming distinct or barely perceptible longitudinal black stripes parallel to each other, crossing

each scale (Fig. 4); sometimes dorsal scales with light-colored posterior edge; the most outer a scale row yellow,

usually with black upper edge; mental and infralabials black-brown; venter yellow in adult specimens, yellowish-

white in juvenile specimens; chin shields, throat, subcaudals usually with black markings; in SYS r000807, each

ventral scale with a middle black spot, formed a longitudinal row along medioventral line.

Coloration in ethanol. Dorsal surfaces blackish-brown, longitudinal stripes indistinct or even not visible;

dorsal scales more or less with pale-colored posterior edges; pale-yellow to whitish beneath, more or less with grey

black spots.

Revision of diagnosis. This species is characterized by the combination of (1) TL 378–462 mm in adult

specimens; (2) TaL 15–20% of TL; (3) rostral concave, slightly large, RW 31–37% of HW; (4) nasal in contact

with first and second supralabials; (5) loreal more than twice as long as deep; (6) loreal entering orbit or not, in

contact with second to fourth or to fifth supralabials; (7) supralabials 7–8, the last one significantly smaller than the

adjacent preceding one; (8) infralabials 7–10; (9) dorsal scale rows 17:17:17; (10) dorsal scales smooth on the

neck, feebly keeled on the middle part of body, rather strongly keeled on posterior part of body, strongly keeled on

the tail; (11) ventrals 141–174; subcaudals 43–60; (12) olive-green to olive-brown above, with distinct or barely

perceptible longitudinal black stripes parallel to each other, crossing each scale; yellow beneath, with black

markings in life.

Distribution and habitat. Opisthotropis andersonii has been recorded in Hong Kong and adjacent Shenzhen

City, extending north to Guangzhou City, China, and in northern and central Vietnam (David et al. 2011). It is a low

mountain dweller, commonly found in montane fast or slow-flowing streams, tiniest trickles covered with bare

rocks, pools and ditches, surrounded by moist subtropical evergreen broadleaved forests.

Remarks. Opisthotropis andersonii was described originally from a single juvenile specimen with TL 240

mm, TaL 45 mm, which was obtained, through the kind mediation of Dr. J. Anderson from the Directors of the City

Hall Museum, Hong Kong, labelled as from Hong Kong, presented to the City Hall Museum by C. Ford

WANG ET AL.

402

(Boulenger 1888). As the City Hall Museum has been closed for a long time and we have been unable to examine

this preserved specimen.

According to the original description of Boulenger (1888), O. andersonii possessed a preocular, two

suboculars and a postocular. Zheng (1992) examined two specimens (BM.217 and 218, collected on 1956)

deposited in the British Museum, London and considered that O. andersonii possessed two preoculars and two

postoculars, no subocular. Obviously, Zheng (1992) treated two suboculars (i.e., pre-subocular and post-subocular)

considered by Boulenger (1888) as lower preocular and lower postocular, respectively. We follow the descriptions

of Zheng (1992). Opisthotropis andersonii is oviparous, probably feeds on semi-aquatic earthworms, and

sometimes does well in captivity on a diet of earthworms (Karsen et al. 1998).

FIGURE 4. General aspect of Opisthotropis andersonii. A: dorsolateral view of the adult female specimen SYS r001020 in

life. B: Ventral view of the specimen SYS r001020. C and D: Adult male specimen SYS r000893 in life. E: Lateral view of

adult female SYS r001382. Photos by JW, JZ, JHY and ZYL.

Opisthotropis shenzhenensis sp. nov.

Figs. 5 and 6a

Holotype. SYS r001018, adult male, collected by Jian Zhao and Run-Lin Li on 15 September, 2014 from Mt.

Wutong (22°34′54.8″N, 114°12′2.7″E; 260 m a.s.l.), Shenzhen City, Guangdong Province, China.

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A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

Paratypes. SYS r000635, an adult male, collected by JHY and RLL on 11 April, 2012 from the same locality

as the holotype. Three adult female specimens: SYS r000636 collected by JHY and RLL on 11 April, 2012 from

the same locality as the holotype; SYS r001021 collected by JZ and RLL on 17 September, 2014 from Sanzhoutian

(22°38′52.16″N, 114°16′32.08″E; 314 m a.s.l.), Shenzhen City; SYS r001032 collected by JZ and RLL on 19

September, 2014 from Mt. Tiantou (22°41′1.75″N, 114°24′17.58″E; 327 m a.s.l.), Shenzhen City; SYS r001145

collected by JW and ZTL on 13 April, 2015 from Mt. Yingping (22°52′40.54″N, 114°9′12.27″E; 155 m a.s.l.),

Dongguan City, Guangdong Province, China.

Etymology. The specific name “shenzhenensis” is derived from the type locality of the new species, Shenzhen

City, Guangdong Province, China. We propose the common name in English “Shenzhen Mountain Stream Snake”

and “Shenzhen Hou Leng She” in Chinese.

Diagnosis. Opisthotropis shenzhenensis sp. nov. differs from other species of Opisthotropis by a combination

of the following characters: (1) TL 328–412 mm; (2) tail short, TaL 16–19% of TL; (3) rostral, narrowed, RW 25–

30% of HW; (4) nasal in contact with first, second and third supralabials; (5) nasal cleft invariably pointing to the

middle of upper edge of second supralabial; (6) loreal 1.3–1.6 times as long as deep; (7) loreal not entering the

orbit, not in contact with second supralabials in all known specimens; (8) supralabials 9–10, the last one smaller

than adjacent preceding one; (9) infralabials 8–10; (10) dorsal scale rows 19:19:19; (11) dorsal scales keeled

throughout, weakly keeled on neck, posteriorly strongly keeled; (12) ventrals 162–179; subcaudals 53–60; (13)

olive-green above in life, each scale with black edge, forming fine mesh pattern; yellow beneath, ventral surface of

head and tail mottled with blackish grey.

Description of holotype. Body cylindrical; TL 407 mm (SVL 337 mm, TaL 70 mm); tail short, TaL 17% of

TL; head small, indistinct from neck; rostral nearly flatted, small, slightly less than twice as broad as deep, RW 25–

30% of HW, barely visible from above; two internasals, crescentic-shaped, in contact with each other medially

behind the rostral, not in contact with loreal, posteriorly in contact with prefrontal; a single prefrontal, in contact

with loreal, preocular and supraocular laterally, with frontal posteriorly; a single frontal, pentagonal, in contact

with supraocular laterally, with two parietals posteriorly; parietals large, in contact with each other medially; nasal

directed dorsolaterally, polygonal, in contact with first, second and third supralabials ventrally, with loreal and

prefrontal posteriorly, with internasal dorsally, with rostral anteriorly; nostril horizontally oval, in the upper part of

nasal; a short vertical cleft below the nostril and dividing nasal into anterior and posterior parts, pointing to the

middle of upper edge of second supralabial; a single loreal, trapezoid, 1.4 times as long as deep, not entering the

orbit, in contact with third and fourth supralabials, but not with second infralabial; a single supraocular; a single

preocular; two postoculars; a single anterior temporal, significantly elongate; two posterior temporals; supralabials

9/9, the last one shorter than the adjacent preceding supralabial; fifth and sixth supralabials entering orbit;

infralabials 9/9, the first one in contact with its fellow behind the mental; two pairs of chin shields; anterior chin-

shields large, in contact with each other medially, and in contact with the first five infralabials on both sides;

posterior chin shields small and separated from each other by small scales; dorsal scale rows 19:19:19; dorsal

scales weakly keeled on neck, posteriorly gradually strongly keeled; the outermost a dorsal scale row smooth at

midbody, those following keeled; V 172; cloacal plate divided; subcaudals 53, paired.

Coloration in life. Olive-green above, each scale with black edge, forming a fine mesh pattern; the outer

second and third scale rows with yellow spots, upper part of the most outer scale row dark green, lower part yellow;

yellow beneath, ventral surface of head and tail mottled with blackish-grey.

Coloration in ethanol. Greenish-black above, barely perceptible black edge on each scale; yellowish beneath.

Va ri at io n . Measurements, scalation and body proportions of type series of Opisthotropis shenzhenensis

sp.nov. are given in Table 5. In specimen SYS r001032, anterior chin-shields in contact with first six infralabials in

both sides. In the SYS r001021, only fifth supralabial entering the orbit in both sides. In the SYS r001032 and

1145, fifth and six supralabials entering the orbit in left side, fifth entering the orbit in right side.

Comparisons. Opisthotropis shenzhenensis sp. nov. is mostly similar to O. andersonii and O. maxwelli in

habitus, color and pattern. In our phylogenetic tree, the new species is the sister taxon of O. andersonii, from which

it differs in having (1) DSR 19:19:19 vs. 17:17:17; (2) dorsal scales keeled throughout, weakly keeled on neck,

posteriorly strongly keeled vs. smooth on the anterior neck, posteriorly feebly keeled, rather strongly keeled on

posterior part of body; (3) rostral small, RW 25–30% of HW vs. rostral slightly large, RW 31–37% of HW; (4)

nasal in contact with first, second and third supralabials vs. only with first and second, not in contact with third; (5)

loreal 1.3–1.6 times as long as deep vs. 2.1–2.9 times as long as deep; (6) loreal not in contact with second

WANG ET AL.

404

supralabial vs. loreal in contact with second supralabial. Opisthotropis shenzhenensis sp. nov differs from O.

maxwelli in having (1) DSR 19:19:19 vs. 17:17:17; (2) tail short, TaL 16–19% of TL vs. TaL 20–23% of TL; (3)

the last supralabial smaller than adjacent preceding one vs. the last supralabial is the longest one; (4) dorsal scales

keeled throughout, weakly keeled on neck, posteriorly strongly keeled vs. smooth on the anterior neck, feebly

keeled posteriorly, rather strongly keeled on posterior part of body; (5) nasal cleft invariably pointing to the middle

of upper edge of second supralabial vs. invariable pointing to the apex angle of first supralabial; (6) nasal in contact

with first, second and third supralabials vs. in contact only with first and second, but not with third. See Fig. 6.

FIGURE 5. General aspect and scalation of head of Opisthotropis shenzhenensis sp. nov.. A: Dorsal view of the holotype SYS

r001018. B: Ventral view of the holotype. C: Dorsolateral view of head and anterior neck of the holotype in life. D: General

aspect of SYS r001145 in life. Photos by JW and JZ.

Opisthotropis shenzhenensis sp. nov., O. andersonii, O. lateralis and O. kuatunensis are sympatric species

from Mt. Wutong, Shenzhen. O. shenzhenensis differs from O. lateralis by having DSR 19:19:19 vs. 17:17:17; a

single preocular vs. two preoculars; uniformly olive-green above with black mesh pattern on dorsal surface of body

and tail vs. brown above, with a distinct longitudinal black stripe on both sides of body. O. shenzhenensis differs

from O. kuatunensis by having 9–10 supralabials, all of which entire vs. 13–16 supralabials, first six to eight entire,

posterior ones divided horizontally; dorsal scales weakly keeled on neck, strongly keeled on body and tail vs.

extremely strongly keeled throughout; olive-green above with black mesh pattern on dorsal body and tail vs. brown

above, with black longitudinal stripes on body and tail.

Opisthotropis shenzhenensis sp. nov. can be easily distinguished from other Chinese congeners as follows: It

differs from O. latouchii in having DSR 19:19:19 vs. 17:17:17; loreal not entering the orbit vs. entering the orbit;

olive-green above with black mesh pattern vs. yellowish green above, with alternate longitudinal black and yellow

stripes. It differs from O. cheni in having DSR 19:19:19 vs. 17:17:17; loreal not entering the orbit vs. entering the

orbit; olive-green above with black mesh pattern vs. dark olive brown above, with yellow crossbars or marks. It

differs from O. laui in having DSR 19:19:19 vs. 25:23:23; olive-green above with black mesh pattern vs. dark olive

above, with pale yellow crossbars. It differs from O. balteata in having DSR 19:19:19 vs. 19:19:17; V 162–179, SC

405

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

53–60 vs.V 190–205, SC 56–86; dorsal scales keeled throughout, weakly keeled on neck, posteriorly strongly

keeled vs. smooth anteriorly, keeled posteriorly; olive-green above with black mesh pattern on dorsal body and tail

vs. yellow or orange-yellow above, with at least 40 pairs of black bands. It differs from O. guangxiensis in having

DSR 19:19:19 vs.17:15:15; dorsal scales keeled vs. smooth; olive-green above with black mesh pattern vs. olive-

brown above, with narrow light yellow crossbars. It differs from O. maculosa in having DSR 19:19:19 vs.15:15:15;

dorsal scales keeled vs. smooth; olive-green above, each scale with black edge vs. olive-brown above, each scale

with single yellow spot. Lastly, it differs from O. jacobi in having DSR 19:19:19 vs.15:15:15; dorsal scales keeled

vs. smooth.

Opisthotropis shenzhenensis differs from the other 10 congeners not occurring in China in having DSR

19:19:19 vs. DSR 17:17:17 in O. daovantieni Orlov, Darevsky & Murphy, 1998 and O. spenceri Smith, 1918,

19:17:17 in O. tamdaoensis Ziegler, David & Vu, 2008, 19:17:15 in O. rugosa (Lidth de Jeude, 1890), 17 dorsal

scale rows at midbody in O. atra Günther, 1872, 19:21:17 in O. durandi Teynié, Lottier, David, Nguyen & Vogel,

2013, 15 dorsal scale rows at midbody in O. kikuzatoi Okada & Takara, 1958; loreal not in contact with internasal

in both sides of head vs. in contact in O. alcalai Brown & Leviton, 1961, O. cucae David, Pham, Nguyen &

Ziegler, 2011, O. typica (Mocquard, 1890), O. atra, O. daovantieni, O. durandi, O. rugosa and O. spenceri; SPL 9–

10 vs. 12–13 in O. alcalai, 7 in O. atra and O. cucae, 6–7 in O. durandi, 12–13 in O. rugosa, 11–12 in O. typica; V

162–179 and SC 53–60 vs. V 195 in O. alcalai, V 191 and SC 44 in O. cucae, V 189–194 and SC 39–47 in O.

daovantieni, SC 88–90 in O. durandi, V 180–198 in O. kikuzatoi, SC 82–95 in O. typica; dorsal scales keeled

throughout vs. smooth throughout in O. alcalai, O. cucae, O. daovantieni, O. durandi, and O. spenceri.

FIGURE 6. Comparisions of head scalation of a: Opisthotropis shenzhenensis sp. nov. (holotype SYS r001018), b: O.

maxwelli (SYS r000841), c and d: O. andersonii (SYS r001020, SYS r001082). Line illustration by ZTL.

Distribution and habitat. Opisthotropis shenzhenensis sp. nov. occurs from the low mountain ranges in the

eastern Shenzhen and southern Dongguan, Guangdong Province, China, including Mt. Wutong, Sanzhoutian and

Mt. Tiantou, and Mt. Yinping (four red dots in Fig. 1). All specimens were collected at night from slow-flowing

streams covered with bare rocks at elevations between 155 and 327 m a.s.l., where we observed a large number of

freshwater snails, aquatic insects, shrimps, crabs, and fishes, and amphibians such as Leptolalax laui and

Paramesotriton hongkongensis occur (Fig. 7).

Remarks. Opisthotropis shenzhenensis, O. andersonii, O. kuatunensis and O. lateralis are sympatric and they

were found frequently in two streams on Mt. Wutong, where a total of 17 specimens were collected between 2012

and 2015, including five of O. kuatunensis, five of O. lateralis (see Appendix), four of O. andersonii and three of

O. shenzhenensis.

WANG ET AL.

406

FIGURE 7. Habitat of Opisthotropis shenzhenensis sp.nov. at the type locality, Mt. Wutong, Shenzhen City, Guangdong

Province, China, which is inhabited also by the other three sympatric species O. andersonii, O. kuatunensis and O. lateralis.

Discussion

The description of O. shenzhenensis brings the total number of species of Opisthotropis to 22, of which 12 occur in

China.

Previous species descriptions were based on the morphology of a few specimens and in some cases, only a

limited number of diagnostic characters were provided. This resulted in considerable challenges for accurate

species identification. The integrative taxonomy is making significant progress in addressing these issues (Dayrat

2005; Grismer et al. 2013). In our study, the scalation of O. andersonii is significantly variable among ten

specimens from southern China: loreal entering the orbit, one preocular or lacking preocular in six specimens, i.e.,

SYS r001424 from Hong Kong, SYS r000803, 1017 and 1082 from Mt. Wutong, SYS r000803 from Mt. Yangtai,

Shenzhen, SYS r001382 and 1383 from Mt. Maofeng, Guangzhou (see Table 4); the remaining four specimens,

i.e., SYS r001423 from Hong Kong, SYS r000893 from Dapeng Peninsula, SYS r000607 and 1020 from Mt.

Wutong, Shenzhen, agree completely with previous descriptions of O. andersonii by Boulenger (1888) and Zheng

(1992). However, our phylogenetic analysis indicated that all ten specimens are clustered into one clade with small

genetic divergence (p-distance ≤ 0.044), representing a monophyletic taxon, which is a sister species to the new

species O. shenzhenensis. Therefore, we considered that these ten specimens are classified as O. andersonii based

on the theories and methods of molecular phylogenetics coupled with morphological characters.

Our field surveys uncovered high levels of Opisthotropis species diversity in lowland mountainous streams

from Hong Kong to Shenzhen, southern China. However, the lowland and submontane forests within this area are

being threatened by land use changes, invasive plants, and ecological degradation. Therefore, surveys are needed

urgently in South China to investigate the population status and distribution of this species and we recommend the

species should be listed as Data Deficient in the IUCN Red List of Threatened Species. Moreover, the known

habitats of these species should be protected by the relevant authorities.

407

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

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O. alcalai <HV   " ± " " ""   "

O. andersonii 1R  ±  ± ±    ± ±

O. atra <HV     " " ""   

O. balteata 1R   ± ± ±    ± ±

O. cheni 1R    ± ±    ± ±

O. cucae <HV     ±     

O. daovantieni <HV         ± ±

O. durandi <HV    ± ˛  ±"  ± ±

O. guangxiensis 1R  ±  ± ±    ± ±

O. jacobi 1R    ±     ± ±

O. kikuzatoi 1R   "  " " ""  ± "

O. kuatunensis 1R  ± ± ± "    ± ±

O. laui 1R     ±    ± ±

O. lateralis 1R    ±     ± ±

O. latouchii 1R  ±  ±     ± ±

O. maculosa 1R   ± ± ±    ± ±

O. maxwelli 1R  ±  ±     ± ±

O. rugosa <HV   ± ± ±    ± ±

O. shenzhenensisVSQRY 1R    ± ±    ± ±

O. spenceri <HV  ±  ± "     

O. tamdaoensis 1R    ± ±     

O. typica <HV  ± ± ± ±  ""  ± ±

VPRRWKZHHNO\NHHOHGGLVWLQFWO\NHHOHGVWURQJO\NHHOHGNHHOLQJDWPLGERG\

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Lastly, we present below a new key to the 12 species of Opisthotropis occurring in China, modified from Zhao

(2006) and Yang et al. (2011). Measurements and morphological data of O. latouchii (n=43) and O. cheni (n=11)

from examined specimens.

1 Dorsal scales in 19 or more rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

- Dorsal scales in 15 or 17 rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5

2 Dorsal scales in 19 rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3

- Dorsal scales in 23 rows at midbody . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..O. laui

3 Dorsal scale rows 19:19:17, ventrals 190–205, body and tail with many pairs of transverse black bands . . . . . . . . . O. balteata

- Dorsal scale rows 19:19:19, ventrals less than 190; dorsal body without transverse black bands . . . . . . . . . . . . . . . . . . . . . . . .4

4 Supralabials 13–16, posterior several ones divided horizontally; dorsal scale strongly keeled throughout; brown above, with

black longitudinal stripes on body and tail . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. kuatunensis

- Supralabials 9–10, not divided horizontally; dorsal scales weakly keeled on neck, strongly keeled on body and tail; olive-green

above, with fine black mesh pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..O. shenzhenensis sp. nov.

5 Dorsal scales in 15 rows at midbody. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6

- Dorsal scales in 17 rows at midbody. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8

6 Dorsal scale rows 17:15:15 body with narrow pale yellow crossbars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. guangxiensis

- Dorsal scale rows 15:15:15; body without pale yellow crossbars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

7 Olive-brown above, with a single yellow spot on each scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. maculosa

- Uniformly dark above, dorsal scales pale-edged posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. jacobi

8 Presence of a lateral black stripe separating abruptly ventral pale color from dorsal dark color, loreal not entering orbit . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. lateralis

- Absence of same lateral black stripe, loreal entering orbit or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

9 Loreal not entering orbit, 1.4–1.7 times as long as deep; the last supralabial longest . . . . . . . . . . . . . . . . . . . . . . . . O. maxwelli

- Loreal entering orbit or not, more than 1.7 times as long as deep; the last supralabial smaller than the adjacent preceding one .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

10 Loreal 2.1–2.9 times as long as deep, in contact with second supralabials; tail length 15–20% of total length; dorsal surface

without yellow longitudinal stripes and crossbars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. andersonii

- Loreal 1.7–2.6 times as long as deep, not in contact with second supralabial; tail length 20–23% of total length; dorsal surface

with yellow longitudinal stripes or crossbars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

11 Yellowish-green above, alternating longitudinal black and yellow stripes, crossing each scale . . . . . . . . . . . . . . . . O. latouchii

- Olive-brown above, with yellow crossbars or yellow marks edged with black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. cheni

Acknowledgements

We thank Jian-Zhao, Hai-Long He and Run-Lin Li from the Museum of Biology Sun Yat-sen University for their

help in the fieldwork. This work was supported by the Project of Wildlife Investigation of Shenzhen and the

specimen platform of China, teaching specimen sub-platform (http://mnh.scu.edu.cn/) to Y.Y. Wang.

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APPENDIX. List of further specimens examined.

Country Location Voucher Number

Opisthotropis cheni (n=11)

China Guangdong Province: Ruyuan County: Nanling Nature

Reserve

SYS r000007, SYS r000010, SYS r000040, SYS

r000045, SYS r000050

China Guangdong Province: Tianjingshan Forestry Station SYS r001293, SYS r001294, SYS r001295;

China Guangdong Province:Lianzhou City: Mt. Dadong: SYS r000934

China Guangdong Province: Yingde City: Shimentai Nature

Reserve

SYS r001422

China Hunan Province: Yizhang County: Mt. Mangshan SYS r000369

Opisthotropis kuatunensis (n = 11)

China Guangdong Province: Huidong County: Baipenzhu

Nature Reserve

SYS r000093; SYS r000094

China Guangdong Province: Shenzhen City: Mt. Wutong SYS r000606, SYS r000608, SYS r000615 and SYS

r001081

China Jiangxi Province: Yongxin County: Qixiling Nature

Resewrve

SYS r000998

China Jiangxi Province: Guangfeng County: Tongboshan

Nature Reserve

SYS r000737, SYS r000745

......continued on the next page

411

A NEW SPECIES OF OPISTHOTROPIS FROM CHINA

APPENDIX 1. (Continued)

Country Location Voucher Number

China Fujian Province: Shanghang County: Wulong Village SYS r001008

Unknown locality SYS r000576

Opisthotropis latouchii (n = 43)

China Guangdong Province: Ruyuan County: Nanling Nature

Reserve

SYS r000008 and SYS r000049

China Guangdong Province: Ruyuan County SYS r000753

China Guangdong Province: Yingde City: Shimentai Nature

Reserve

SYS r000764

China Jiangxi Province: Mt. Jinggang SYS r000296, SYS r000300, SYS r000309, SYS

r000358, SYS r000359, SYS r000421, SYS r000422,

SYS r000423, SYS r001253, SYS r001255, SYS

0001298, SYS r001300, SYS r001303, SYS r001304,

SYS r001305 and SYS r001306

China Jiangxi Province: Jing’an County: Jiulingshan Nature

Reserve

SYS r000987, SYS r000991 and SYS r000993

China Jiangxi Province: Chongyi County: Qiy unshan Nature

Reserve

SYS r000961

China Jiangxi Province: Anfu County: Mt. Wugong SYS r000979

China Jiangxi Province: Xinfeng County: Mt. Jinpeng SYS r001370

China Jiangxi Province: Anyuan County: Sanbaishan SYS r001004

China Jiangxi Province: Guangfeng County: Tongboshan

Nature Reserve

SYS r000776

China Hunan Province: Yanling County: Taoyuandong Nature

Reserve

SYS r000823, SYS r000824 and SYS r000825

China Hunan Provence:Yizhang County: Mt. Mangshan SYS r000368 and SYS r000371

China Hunan Provence: Hengshan County: Mt. Hengshan SYS r001322

China Fujian Province: Wuyishan City: Guadun (=Kuatun)

Village: Sangang Village

SYS r000670

China Fujian Province: Wuyishan City: Sangang Village SYS r000671, SYS r000677

China Fujian Province:Shaowu City: Longhu Forestry Station SYS r001287

China Guangxi Zhuang Autonomous Region: Jinxiu County:

Dayaoshan Nature Reserve

SYS r000843

China Guangxi Zhuang Autonomous Region: Lingui County:

Anjiangping

SYS r000853

China Guizhou Province: Leishan County: Mt. Leigong SYS r000857 and SYS r000866

Opisthotropis lateralis (n = 11)

China Guangdong Province: Xinyi County: Dawuling Nature

Reserve

SYS r000092

China Guangdong Province: Shenzhen City: Mt. Wutong SYS r000611, SYS r000612, SYS r000616, SYS

r001019, SYS r001080 and SYS r001389

China Guangdong Province: Fengkai County: Heishiding

Nature Reserve

SYS r000951

China Guangdong Province: Zhaoqing City: Mt. Dinghu SYS r001149

China Guangxi Zhuang Autonomous Region: Jingxi County:

Xinling Village

SYS r000262

China Hainan Province: Dazhipo SYS r000361

......continued on the next page

WANG ET AL.

412

APPENDIX 1. (Continued)

Country Location Voucher Number

Opisthotropis laui (n = 4)

China Guangdong Province: Taishan City: Shangchuan Island SYS r001161, SYS r001169, SYS r001170 and SYS

r001171

Opisthotropis guangxiensis (n = 3)

China Guangdong Province: Xinyi County, Dawuling Nature

Reserve

SYS r001448 and SYS r001449

China Guangxi Zhuang Autonomous Region: Jinxiu County:

Mt. Dayao

SYS r001477

Opisthotropis maculosa (n = 7)

China Guangxi Zhuang Autonomous Region: Shangsi County:

Shiwanshan Nature Reserve

SYS r000393, SYS r000394 and SYS r000537

China Guangdong Province: Pubei County: Wuhuangling

Forest Park

SYS r000538

China Guangdong Province: Fengkai County: Heishiding

Nature Reserve

SYS r000720, SYS r000946 and SYS r001350

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Jun 2020
ZOOTAXA

A new species of natricid snake, Smithophis linearis sp. nov., is described on the basis of a single recently collected specimen from Yingjiang County, Yunnan Province, People's Republic of China, and three historical specimens from Yunnan and from northeastern Myanmar. The new species is assigned to the genus Smithophis on the basis of its single internasal and single prefrontal shields, and on the basis of the results of phylogenetic analysis of mitochondrial cytochrome b DNA sequence data. The new species differs from its congeners in having the following combination of characters: temporal shields present, six or more circumorbital scales, and a distinctive colour pattern comprising regular, narrow, longitudinal dark and pale lines. Morphological and cytochrome b data are consistent with the recognition of Smithophis as distinct from the genus Opisthotropis. A revised key to the identification of the species of Smithophis is provided.

A Large-scale Systematic Framework of Chinese Snakes Based on a Unified Multilocus Marker System

Article

Apr 2020

Notes on the distribution and natural history of the enigmatic and endangered Opisthotropis alcalai (Brown & Leviton, 1961) on Mindanao Island, Philippines

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Jul 2023

Snakes of the World: A Supplement

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Aug 2020

Jeff Boundy

Re-examination of the Chinese record of Opisthotropis maculosa (Squamata, Natricidae), resulting in the first national record of O. haihaensis and description of a new species

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Feb 2020

The taxonomic status of the previous record of Opisthotropis maculosa Stuart & Chuaynkern, 2007 from Guangdong and Guangxi, southern China, is revised based on the comparison of morphological and molecular data collected from the Chinese specimens and the holotype of O. maculosa from Thailand and O. haihaensis Ziegler, Pham, Nguyen, Nguyen, Wang, Wang, Stuart & Le, 2019 from Vietnam. Results reveal that the population from Shiwandashan Nature Reserve in southern Guangxi, China belongs to O. haihaensis, and represents the first national record for China; the populations from western Guangdong and southeastern Guangxi are described as a new species, Opisthotropis hungtai sp. nov. We suggest that O. maculosa should be removed from the Chinese herpetofauna checklist. The new national record of O. haihaensis and the description of the new species bring the total number of Opisthotropis to 13 in China.

On the second specimen of Opisthotropis rugosus (van Lidth de Jeude, 1890) (Colubridae, Natricinae)

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Oct 2009

Opistothropis rugosus (van Lith de Jeude, 1890), hitherto known from the unique holorype, is here reported after over a century, ca. 200 km north-west of the type locality. This specimen is similar to O. typicus (Mocquard, 1890) (type locality: Gunung Kinabalu, Sabah, Malaysia). Additional specimens of O. typicus, reported for the first time from Kalimantan Tengah (Borneo), south of the equator, is reported here.

A new species of the genus Opisthotropis Günther, 1872 (Squamata: Natricidae) from the highlands of Kon Tum Province, Vietnam

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Feb 2011

A new species of the snake genus Opisthotropis Günther, 1872, Opisthotropis cucae sp. nov., is described from Kon Tum Plateau in central Vietnam. It is distinguished from any other Opisthotropis species by the combination of the following characters: (1) dorsal scales entirely smooth, in 23-19-19 rows; (2) 191 ventrals; (3) 1 loreal, in contact with internasal; (4) 7 supralabials, 5th entering orbit; (4) dorsum uniformly greyish-brown without bands or crossbars, with an irregular separation between dark dorsum and venter, greyish-yellow speckled with dark grey. An extended comparison with other species of the genus Opisthotropis species is provided together with a key to the known species of the genus. Opisthotropis cucae sp. nov. is the third new mountain stream snake described from Vietnam in the past decade. The grammatical gender of the genus Opisthotropis is discussed; this genus is feminine.

On the taxonomic status of the Thai endemic freshwater snake Parahelicops boonsongi, with the erection of a new genus (Squamata: Natricidae)

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Apr 2015

Parahelicops boonsongi Taylor & Elbel, 1958 is known from only three specimens from Thailand. It has been placed either in the genus Parahelicops Bourret, 1934, along with Parahelicops annamensis Bourret, 1934, or in the genus Opisthotropis Günther, 1872. We compared its morphological characters with those of P. annamensis and with three other relevant genera, Opisthotropis, Pararhabdophis Bourret, 1934, and Paratapinophis Angel, 1929. Parahelicops boonsongi is phenotypically distinct from Parahelicops annamensis, Opisthotropis, and all other natricine genera. We consequently erect a new genus, Isanophis gen. nov., to accommodate Parahelicops boonsongi.

Revision of the diagnostic characters of Opisthotropis maculosa with notes on its distribution and variation, and a key to the genus Opisthotropis (Squamata: Natricidae)

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Mar 2011
ZOOTAXA

Opisthotropis maculosa was described based on a single specimen collected from northeastern Thailand in 2007. We collected three specimens from Guangxi and Guangdong, China. These additional specimens provide the first information on morphological variation in this species, including sex-related differences. Based on the new information, we revise the diagnostic characteristics of this species and provide a key to the currently recognized species of Opisthotropis.

New records of reptiles from northern Vietnam

Article

Jan 2014

Band 47 (2) Dezember 2014 Herausgegeben im Selbstverlag des Naturwissenschaftlichen Vereins zu Bremen Abbildung auf dem Umschlag Lacerta bilineata. Foto: Kerstin Elbing © 2014 NWV, Bremen Alle Rechte vorbehalten. Nachdruck, auch auszugsweise, sowie photomechanische, digitale und andere Vervielfältigungen bedürfen der schriftlichen Genehmigung des Naturwissenschaftlichen Vereins zu Bremen. Für den sachlichen Inhalt sind die Autoren verantwortlich. Naturwissenschaftlicher Verein zu Bremen gegr. 1864 Mitglied der Wittheit zu Bremen Postanschrift: Naturwissenschaftlicher Verein zu Bremen c/o Übersee-Museum Bahnhofsplatz 13, 28195 Bremen E-Mail: info@nwv-bremen.de Internet: www.nwv-bremen.de

A New Opisthotropis (Serpentes: Colubridae: Natricinae) from Northeastern Thailand

Article

Jan 2007
Curr Herpetol

A new species of the aquatic natricine colubrid snake genus Opisthotropis is described based on a single specimen from Nong Khai Province, northeastern Thailand. The new species is distinguished from other Opisthotropis by the combination of having smooth scales on the body and tail, 15 dorsal scale rows throughout the body, the fourth supralabial in contact with the eye, the posterior pair of chin shields longer than the anterior pair, and a glossy black dorsum with yellow spots. © 2007, The Herpetological Society of Japan. All rights reserved.

Seven new reptiles from Fukien Province, China

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