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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

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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.
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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Accepted by T. Nguyen: 7 Feb. 2017; published: 29 Mar. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2017 Magnolia Press
Zootaxa 4247 (4): 391
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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
2
, ZU-YAO LIU
1
, ZHI-TONG LYU
1
, JIAN WANG
1
, LIN LUO
2
,
YAN-JUN SUN
3
& YAN-WU ZHANG
2
1
State Key Laboratory of Biocontrol / The Museum of Biology, School of Life Sciences, Sun Yat-sen University, Guangzhou 510275,
P.R. China
2
Shenzhen Administrative Bureau for Wildlife and Forest Plants Protection, Shenzhen 518048, P.R. China
3
Shenzhen Park Service, Shenzhen 518040, P.R. China
4
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.
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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;
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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
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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.
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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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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              
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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
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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.
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A NEW SPECIES OF OPISTHOTROPIS FROM CHINA
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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
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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
th
vertebral scale in SYS r001423, on 13
th
in SYS r001424, 15
th
in SYS r000803 and 1382, 17
th
in SYS r001020, 18
th
in SYS r001082, 21
st
in SYS r000607, 30
th
in SYS r001017, 46
th
in SYS r001383, 58
th
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
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(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
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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
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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.
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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.
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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. shenzhenensisVS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
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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
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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
... Most later authors followed Pope (1935) and considered Trimerodytes to be a junior synonym of Opisthotropis and its type species, T. balteatus, has been assigned to Opisthotropis (Brown and Leviton, 1961;Nguyen et al., 2009;Smith, 1943;Zhao, 2006;Zhao et al., 1998;Zheng, 1992). Although there have been recent phylogenetic studies on Opisthotropis, none of them included O. balteata in their analyses (Ren et al., 2017;Wang et al., 2017b;Ziegler et al., 2017;. Therefore, the generic assignment of O. balteata has not been confirmed by molecular data. ...
... mitochondrial gene cytochrome b (cyt b) was targeted and amplified via the polymerase chain reaction (PCR), using the following primer pairs: L14919 (5'-AACCACCGTTGTTATTCAACT-3')/L14910 (5'-GACCTGTGATMTGAAAACCAYCGTTGT-3'), and H16064 (5'-CTTTGGTTTACAAGAACAATGCT TTA-3') (Burbrink et al., 2000;Guo et al., 2012;Wang et al., 2017b), PCR protocols as described by Ren et al. (2017). The PCR products were purified and sequenced in both directions using an ABI 3730xL sequencer by Sangon Biotech Co., Ltd (Chengdu, China). ...
... Morphological and ecological data of related taxa were based on specimens examined (Appendix 1) and from the literature (Brown and Leviton, 1961;Chuaynkern et al., 2014;David et al., 2011;Günther, 1872;Huang et al., 1990;Iskandar and Kamsi, 2009;Le et al., 2015;Murphy et al., 2008;Noonloy et al., 2018;Okada and Takara, 1958;Orlov et al., 1998;Ota, 2004;Pauwels et al., 2009;Pope, 1935;Rao and Yang, 1998;Rasmussen, 1982;Ren et al., 2017;Rossman and Eberle, 1977;Smith, 1943;Stuart and Heatwole, 2008;Teynié et al., 2014;Toyama, 1983;Vogel et al., 2004;Wang et al., 2017aWang et al., , 2017bWu et al., 1985;Yang et al., 2011Yang et al., , 2013Zhao, 2006;Zhao et al., 1998;Zheng, 1992;Ziegler et al., 2017Ziegler et al., , 2018. ...
Article
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The single prefrontal configuration has historically been used as an important diagnostic character for many natricine taxa. For example, the genus Trimerodytes Cope, 1895 was long been regarded as a junior synonym of Opisthotropis Günther, 1872 for their similar prefrontal configurations and the type species, T. balteatus Cope, 1895, has been assigned to the genus Opisthotropis. However, as the number and arrangement of prefrontal vary frequently both at species and generic level, it is questionable whether the synonymization of Trimerodytes reflects their evolutionary relationships. On the basis of recently collected specimens of O. balteata, the generic status of the species was assessed using both molecular and morphological data. Opisthotropis was recovered as polyphyletic with reference to O. balteata, because O. balteata is nested within the genus Sinonatrix Rossman and Eberle, 1977 and is the sister species of the type species of Sinonatrix. Consequently, we herein resurrect the long-overlooked synonym Trimerodytes from Opisthotropis and synonymize the junior generic nomen Sinonatrix with Trimerodytes. In addition, based on morphological similarities between the monotypic genus Paratapinophis Angel, 1929 and Trimerodytes, we doubt about the validity of Paratapinophis. Following taxonomic changes in this work, the taxonomic account of the genus Trimerodytes, updated descriptions of its type species, and diagnostic key to Trimerodytes species are provided.
... Mountain Stream Keelbacks can be found across the mainland of Southeast Asia to Sumatra (Indonesia), the Philippines and the Ryukyu Archipelago of Japan. The genus, which currently consists of 24 species, is still poorly known (Teynié et al., 2013;Ren et al., 2017;Wang et al., 2017;Ziegler et al., 2018). Eight species have been discovered in the past eleven years: O. laui Yang, Sung & Chan, O. shenzhenensis Wang, Guo, Liu, Lyu, Wang, Luo, Sun & Zhang and O. zhaoermii Ren, Wang, David, Pham, Nguyen & Ziegler, O. tamdaoensis Ziegler, David & Vu and O. voquyi Ziegler, David, Ziegler, Pham, Nguyen & Le from Vietnam (Stuart & Chuaynkern, 2007;Ziegler et al., 2008;David et al., 2011;Teyni et al., 2013;Yang et al., 2013;Ren et al., 2017;Wang et al., 2017;Ziegler et al., 2017Ziegler et al., , 2018. ...
... The genus, which currently consists of 24 species, is still poorly known (Teynié et al., 2013;Ren et al., 2017;Wang et al., 2017;Ziegler et al., 2018). Eight species have been discovered in the past eleven years: O. laui Yang, Sung & Chan, O. shenzhenensis Wang, Guo, Liu, Lyu, Wang, Luo, Sun & Zhang and O. zhaoermii Ren, Wang, David, Pham, Nguyen & Ziegler, O. tamdaoensis Ziegler, David & Vu and O. voquyi Ziegler, David, Ziegler, Pham, Nguyen & Le from Vietnam (Stuart & Chuaynkern, 2007;Ziegler et al., 2008;David et al., 2011;Teyni et al., 2013;Yang et al., 2013;Ren et al., 2017;Wang et al., 2017;Ziegler et al., 2017Ziegler et al., , 2018. Nine species of the Mountain Stream Keelbacks have been recorded so far from Vietnam (David et al., 2011, Ziegler et al., 2017Nguyen et al., 2018;Ziegler et al., 2018); most of them are uncommon or at least rarely collected. ...
... Morphological comparisons of the new species with other species of the genus Opisthotropis and Hebius annamensis (formerly Parahelicops annamensis; see Kizirian et al. 2018) were based on data from Stuart & Chuaynkern (2007), David et al. (2011), Teynié et al. (2013, Nguyen et al. (2017), Ren et al. (2017), Wang et al. (2017), andZiegler et al. (2018). ...
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The Yellow-spotted Mountain Stream Keelback Opisthotropis maculosa was originally described based on a single male specimen from northeastern Thailand. Recently, based on morphological data, new records of this species were published initially from southern China and subsequently from northern Vietnam. In this study, we provide the first molecular comparisons between the holotype and other populations in China and Vietnam using the mitochondrial cytochrome b gene and use an integrative taxonomic approach to show that the population from Vietnam represents a distinct taxon. Opisthotropis haihaensis sp. nov. is characterized by a combination of the following characters: internasal not in contact with loreal; prefrontal not touching supraocular; frontal touching preocular; one preocular; one postocular; one anterior temporal; one posterior temporal; eight supralabials, fourth and fifth in contact with eye; 24 maxillary teeth; anterior pair of chin shields longer than posterior pair; 169 ventrals + 2 preventrals); 79 subcaudals, paired; 15 dorsal scale rows at neck, at midbody and before vent; body and tail scales smooth; chin shields yellow with brownish black mottling; body and tail dorsum dark with each a light spot per scale. Phylogenetically, the new species is supported as the sister taxon to “O. maculosa” from China (but separated by approximately 10% uncorrected pairwise sequence divergence) and is distantly related to O. maculosa sensu stricto from Thailand, warranting a taxonomic revision of the maculosa-like species. According to our results, O. maculosa should be delisted from the herpetofauna of Vietnam, which currently consists of nine Opisthotropis species. Five species, O. cucae, O. daovantieni, O. haihaensis, O. tamdaoensis, and O. voquyi, are endemic to the country.
... Nowadays, only two complete mitochondrial genomes of Opisthotropis species are published (Wang et al. 2019;Zhang et al. 2019). The genetic diversity of this species is still unclear in the Natricidae due to limited genetic studies (Wang et al. 2017). In this study, we sequenced the mitogenome of O. kuatunensis using next-generation sequencing to determine its mitogenome structure and systematic relationship to other Natricidae. ...
... The p-distances between O. kuatunensis and other Opisthotropis species were both more than 7% based on the 13 PCGs via MEGA 5.05. This mitogenome obtained in this study enriches the genomic resources available and fully unravel the phylogenetic relationship among Opisthotropis species, which is was in accordance with previous works using morphology and DNA markers (Wang et al. 2017(Wang et al. , 2019Zhang et al. 2019). Further studies on representatives of this family are needed and will be useful for species identification and conservation. ...
Article
Full-text available
Opisthotropis kuatunensis is classified in the family Natricidae and is widespread in southern China. In this study, we sequenced and analyzed the circular mitochondrial genome of O. kuatunensis from the Fujian Province, China. The complete mitogenome is 17,279 bp in length, and includes 13 protein-coding genes, 2 ribosomal RNAs, 22 transfer RNA, 1 non-coding region of an L-strand replication origin and 2 control regions (D-loop1 and D-loop2). Phylogenetic analysis based on the complete mitochondrial genome supported Opisthotropis as monophyletic and sister to Nerodia and fully resolved O. kuatunensis on a branch with O. latouchii. This study contributes to the systematics, phylogeny and taxonomy of the Natricidae.
... Ventral scales were counted following Dowling (1951). We compared the characters of other Sinonatrix and Opisthotropis species, which were obtained from previous research (Zhao et al., 1998;Zhao, 2006;Rao & Yang, 1998;Yang et al., 2013;Wang et al., 2017;Ziegler et al., 2018). The skull of YBU 15296 was scanned using computerized tomography (CT) with a high-resolution X-ray scanner (Quantum GX Micro-CT Imaging System, PerkinElmer, Waltham, USA) at the Chengdu Institute of Biology (CIB), Chinese Academy of Sciences (CAS). ...
... differs from most species of Opisthotropis by having 19 rows of dorsal scales at mid-body (vs. 15, 17, or 23 rows Wang, Guo, Liu, Lyu, Wang, Luo, Sun & Zhang, O. spenceri Smith, O. tamdaoensis Ziegler, David & Vu, O. voquyi Ziegler, David, Ziegler, Pham, Nguyen & Le, and O. zhaoermii Ren, Wang, Jiang, Guo & Li (Zhao, 2006;Yang et al., 2013;Wang et al., 2017;Ren et al., 2017;Ziegler et al., 2018). Sinonatrix yapingi sp. ...
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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
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