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Asian Herpetological Research 2020, 11(2): 95–107
DOI: 10.16373/j.cnki.ahr.190068
ORIGINAL
ARTICLE
1. Introduction
The genus Rhabdophis is a group of widely distributed snakes
with a unique defensive system that relies on nuchal gla nds
(Mori et al., 2012). In 1960, Malna te defined this genus based
on morphological characters, i.e. hemipenes with sulcus
spermaticus divided; last two maxillary teeth strongly enlarged,
recurved and usually preceded by a diastema ; internasals broad
anteriorly, nostrils la teral; apical pits present or a bsent; vertebral
gla nds present in several species (M a l na t e, 19 60 ) .
Currently, 27 species are recognized a nd are distributed
in eastern and southern Asia (Zhu et al., 2014; Takeuchi et
al., 2018). Two new species were discovered in the past five
years, Rhabdophis akraios (Doria et al., 2013) and Rhabdophis
guangdongensis (Zhu et al., 2014 ).
A New Species of the Genus Rhabdophis Fitzinger, 1843 (Squamata: Colubridae)
in Southwestern Sichuan, China
Yige PIAO1,2#, Zening CHEN1,3#, Yanqing WU4, Shengchao SHI1,3, Hirohiko TAKEUCHI5, Teppei JONO6, Masaya
FUKUDA7, Akira MORI7, Yezhong TANG1, Qin CHEN1* a nd Li DING1*
Keywords
Rhabdophis chiwen sp. nov., morphology,
Natricinae, phylogenetics, taxonomy
1 Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China
2 University of Chinese Academy of Sciences, Beijing 100049, China
3 Key Laboratory of Bio-Resource and Eco-Environment of Ministry of Education, College of Life Sciences, Sichuan University,
Chengdu 610065, Sichuan, China
4 Nanjing Institute of Environmental Sciences, Ministry of Ecology and Environment of China, Nanjing 210042, Jiangsu, China
5 College of Bioresource Sciences, Nihon University, Fujisawa, Kanagawa 2520880, Japan
6 Tropical Biosphere Research Center, University of the Ryukyus, Nishihara, Okinawa 9030213, Japan
7 Department of Zoology, Graduate School of Science, Kyoto University, Sakyo, Kyoto 6068502, Japan
#Both authors contribute equally to this work.
*Corresponding authors: Dr. Li DING, from Chengdu Institute of
Biology (CIB), Chinese Academy of Sciences (CAS), with his research
focusing on biogeography, systematics, a nd taxonomy of reptiles (snakes);
Dr. Qin CHEN, from CIB, CAS, China, with his research focusing on
behaviour and electrophysiology of infra red organs of Crotalidae,
acoustic communication of Anura.
E-mail: dingli@cib.ac.cn (Ding L.); chenqin@cib.ac.cn (Chen Q.)
Received: 19 December 2019 Accepted: 20 April 2020
Abstract The genus Rhabdo phis is a group of widely
distributed snakes with more tha n 20 species. Recent field
surveys uncovered a species in southwestern China, which
has long been considered as R. pentasu pralabialis. Combined
molecular and morphological analyses revealed it as a new
species Rhabdophis chiwen sp. nov. Based on 12 specimens,
this new species is distinguished by the following
characters: 1) dorsal body saddlebrown, dorsal scales
typically with black margins forming spots and stripes, the
margin of the outer row forming two fa int dorsolateral
bla ck c ross-ba rs al on gs id e bod y; 2 ) ve nt ral sca les 151–159,
the outer margin of ventral scales and several lateral
rows of dorsal scales forming ventrolateral longitudinal
brownish-red coloration, with fa int black spots in the
middle of ventral scales; 3) a black oblique stripe present
below eyes, of ten with a black spot between the 2nd a nd
3rd suprala bial a nd a black stripe on the 5th suprala bial; 4)
eyes dark khaki, pupils black; 5) infrala bials usually 7, the
first four in contact with anterior chin-shields; 6) temporal
scales 1+1; 7) dorsal scales in 15 rows, feebly keeled except
the outer 1–2 rows; 8) anal scale divided; subcaudals 45–59;
9) preocula r 1 a nd postocula rs 3 (occasionally 2); 10) body
medium-sized (snout-vent length: adult males 404–431 mm,
adult females 409–476 mm); 11) tail moderate (tail length/
total length in adult males 0.205–0.238, in adult females
0.172–0.193). With the discovery of this new species, the
total number of species in genus Rhabdo phis is 28 with 12th
species kno wn to occur in China.
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Eleven species of Rhabdo phis are now known to occur
in China (Zhu et al. , 2014), which a re R. adleri ZH AO , 1997 ,
R. guangdongensis Zhu, Wa ng, Takeuchi a nd Zhao, 2014, R.
himala yanus Günther, 1864, R. lateralis Berthold, 1859, R. leonardi
Wall, 1923, R. nigrocinctus Blyth, 1855, R. nuchalis Bou lenger, 1891,
R. pentasu pr alabialis Jia ng and Zhao, 1983, R. subminiatus Schlegel,
1837 , R. f ormosanus Ma ki , 1931 a nd R. swinhonis Günther , 1868.
During several field work in the southwestern part of Sichuan
Province, China, we collected a number of samples which were
originally referred to as R. pentasu pr alabialis. Nevertheless, those
snakes can be distinguished from R. pentasu pralabialis based on
morphological characters. Moreover, the phylogenetic analysis
based on the mitochondrial cytochrome b (c yt b) and nuclear
oocyte ma tura tion factor (cmos) gene sequences revealed tha t
those sa mples a re different from R. pentasu pralabialis and other
congeners in China. erefore, we propose tha t those specimens
be a new species in the genus Rhabdophis.
2. Materials and Methods
2.1. Specim ens samplin g Specimens were collected from
the Xingou Village (XG, 10 specimens) in Tianqua n County
and Jiguan Mountain ( JG, 2 specimens) in Chongzhou City,
Sichuan Province, China (Table 1). Fresh liver or muscle tissues
were taken and immedia tely preserved in 95% ethanol and
stored at –20 ℃. The specimens were dehydra ted in absolute
ethanol a nd then la ter transferred to and stored in 70% etha nol.
In addition, a nother 10 samples of R . leonardi (4 specimens), R.
nuchalis (4 specimens) and R. pentasu pralabialis (2 specimens)
that represent 3 sympa tric species with relatively close niche in
Sichuan Province, were used for molecula r analyses.
2.2. Molecular phylogenetic analysis Genomic DNA was
extracted from the collected tissue samples in Chengdu Institute
of Biology (CIB). PCR amplifications were performed in 25
μl reactions (12.5 μl I-5™ 2×High-Fidelity Master Mix, 10 μl
ddH2O, 1 μl F-primers, 1 μl R-primers, 0.5 μl DNA template)
using the following cycling conditions: initial denatura tion
for 2 min a t 95°C, followed by 35 cycles: denaturation at 94
°C for 40 s, annealing at different tempera tures (48.5 ℃ for c yt
b or 56 ℃ for cmos) for 25 s, elonga tion a t 72 °C for 15 s, and
then finalized with elongation step of 2 min a t 72 °C, with the
PTC-100 thermal cycler (BioRad, USA). Purified PCR products
were sequenced using the same PCR primers. Sequencing was
completed by Beijing Qingke New Industry Biotechnology
Co., Ltd. Sequences for compa rison of a va ila ble species were
downloaded from GenBank. Am phiesma stolatum and N atrix
natrix were used as outgroups.
e c yt b and cmos sequences were combined into one dataset
to build gene trees. All sequences were aligned with other
retrieved sequences in the same gene loci by using software
MEGA 7 (Kumar et al., 2016). Raw trace files were edited in
Geneious 7 (Biomatters Limited, New Zealand). Partition
finder 2.1.1 under BIC identified the optimal models of sequence
evolution for each partition (Lanfear et al., 2 012 ) . Phylogenetic
relationships derived from the combined gene fragments were
performed based on Bayesian Inference (BI) by using MrBayes
3.2 (Ronquist et al., 2 012 ) . We ran our analysis for 20 million
generations with the chains, sampling every 1000 generations.
e average standard deviation of split frequencies (ASDSF <
0.01). e first 1000 trees (of 20,000) were discarded as the burn-
in. A 50 majority-rule with compa tible groups consensus was
taken from the remaining trees and posterior probabilities
(pp) of 0.95 or above were considered significant. We also
performed maximum-likelihood (ML) analysis by using the
program RaxML v8 (Stamatakis, 2014) a nd IQ tree (Nguyen et
al., 2015 ) . Reliability of the ML tree was assessed by calcula ting
bootstrap probability (BP) with 1000 replica tions. Additionally,
Table 1 Morphological characters of Rhabdophis chiwen sp. nov.
No. SVL Tal SPO PRO PTO LR TEM SL IL VEN DSR SC Sex Locality
Holotype CIB116092 422 118 1 1 L:2 R:3 1 1+1 5 L:7R:6 152 15-15-15 56 Male JG
Para type CIB116093 431 131 1 1 3 1 1+1 5 7 155 15-15-15 57 Male JG
Para type CIB116094 411 113 1 1 3 1 1+1 5 7 154 15-15-15 55 Male XG
Para type CIB116095 461 96 1 1 L:3 R:2 1 1+1 5 L:6R:7 157 15-15-15 45 Female XG
CIBDL1807132 427 128 1 1 3 1 1+1 5 L:8R:7 154 15-15-15 58 Male XG
CIBDL1807133 409 128 1 1 3 1 1+1 5 7 157 15-15-15 58 Male XG
CIBDL1807134 411 98 1 1 L:3 R:2 1 1+1 5 L:7R:6 159 15-15-15 49 Female XG
CIBDL1807135 476 107 1 1 3 1 1+1 5 7 156 15-15-15 46 Female XG
CIBDL1807137 409 96 1 1 3 1 1+1 5 L:7R:6 159 15-15-15 52 Female XG
CIBDL1807138 427 122 1 L:2 R:1 L:1 R:3 1 1+1 5 7 152 15-15-15 59 Male XG
CIBDL1807139 404 104 1 1 3 1 1+1 5 7 151 15-15-15 55 Male XG
CIBDL18071311 422 91 1 1 3 1 1+1 5 7 158 15-15-15 47 Female XG
Note: See Material a nd Methods section for the abbreviations.
Yige PIAO et al.
A New Species of Rhabdophis
in Southwestern China
No. 2 97
the uncorrected c yt b and cmos p-dista nce ma trix was com pared
using MEGA 7.0 (Kumar et al., 2016).
2.3. Morphological analysis Twelve adult snakes (7 males and
5 females) were exa mined. Informa tion on some morphological
cha racters of known species were obta ined from litera ture
(Boie, 1827 ; Boulenger, 1896, 1900, 1906; Bourret, 1935; Cantor,
1839; D as, 2010 ; David a nd V o g el, 2010; d e L a ng a nd V og e l ,
2006; Doria et al., 2013; Duméril et al. , 18 54; G ün th er , 1858, 1864 ;
Leviton, 1970; Smith, 1943; Stuebing and Lian, 2002; Tweedie,
1953; Z h ao, 1997 ; Z h ao et al., 1998; Zhao a nd Adler, 1993; Zhao
and Jiang, 1981) and are shown in Table 2. Color description
was done according to wiki color-coding.
Terminology for morphological measurements and
descriptions is as follows: body and tail length were measured
with a tape ruler to the nearest 1 mm; total length (TL), from
the tip of snout to the tip of tail; snout-vent length (SVL),
from the tip of snout to posterior margin of cloaca; tail length
(TaL), from posterior margin of cloaca to the tip of tail. Other
measurements were conducted with a digital caliper to the
nearest 0.1 mm: head length (HL), from the snout tip to the
posterior margin of the mandible; head width (HW) was
measured at the widest part of the head on posterior side; the
eye horizontal diameter (ED), the grea test dia meter of the orbit.
Biometric measures were performed exclusively on the right
eye. Ratio of tail length to total length (TaL/TL) was recorded.
e dorsal scale rows (DSR) were counted at one head length
behind head, at mid-body, and at one head length before vent;
the number of ventral scales ( VEN) was counted according to
the method proposed by Dowling (Dowling, 1951), half ventral
was counted as one. For subcaudals (SC), first scale under the
tail meeting its opposite was regarded as the first subcaudal
scale, and the unpaired terminal scute was not included in the
number of subcaudals. Paired scales on head were counted
on both sides of the head a nd presented in left/right order.
Species DSR Outer dorsal scale row VEN SC PRO PTO MT▲
R.chiwen sp. nov. 15–15–15 smooth 151–159 45–59 1–2 2–3 no data
R. pentasupralabialis 15–15–15 smooth 135–162 43–64 1 2–3 18
R. adleri 19–19–17 feebly keeled 150–164 73–88 1–2 3–4 27(25+2)
R. akraios 19–19–17 keeled 167–184 44–66 1 3 19(17+2)
R. angeli 16(15)–15–15 smooth 117–126 39–46 1 3 22–23
R. auriculata 17–17–15 strongly keeled 157–162 76–91 1–2 3 27–32
R. barbouri 19 at midbody strongly keeled 148–166 96–101 2 3 no da ta
R. callichroma 19 at midbody keeled 152–159 79–86 1–2 3 27–35
R. callistus 21 at midbody strongly keeled 156 76 1 4 no da ta
R. ceylonensis 19 at midbody strongly keeled 137–143 40–54 1 3 24–26(22–24+2)
R. chrysargoides 21 a t midbody strongly keeled 154–161 64–79 1 3 24(22+2)
R. chrysargos 19 at midbody strongly keeled 143–175 60–93 1–2 3 27–35
R. conspicillatus 19 at midbody keeled or smooth 138–147 40–53 1 3 no data
R. flaviceps 19 at midbody keeled 120–138 49–60 1 3–4 22–23
R. guangdongensis 15–15–15 smooth 126 39 1 2 20
R. himalayanus 19–19–17 feebly keeled 165–171 82–88 2 3 26(24+2)
R. leonardi 18(17)–17–15 smooth 149–159 43–62 1 2–3 19(17+2)
R. lineatus 19 a t midbody strongly keeled 132–142 66–71 2 3 18(16+2)
R. murudensis 19–19–15(17) feebly keeled 176–185 63–97 1 3 23(21+2)
R. nigrocinctus 19–19–17 keeled or smooth 150–170 80–97 1 3–4 28(26+2)
R. nuchalis 15–15–15 smooth 144–169 35–65 1 3 18–22
R. plumbicolor 23–27 at midbody strongly keeled 144–160 27–45 2 3–4 no data
R. rhodomelas 19 at midbody strongly keeled 126–136 41–56 1–2 3–4 no da ta
R. rudis 23–23–19 strongly keeled 123–155 44–60 3 3–4 13–20(11–18+2)
R. spilogaster 19 at midbody feebly keeled 148–156 75–92 1–2 3–4 15
R. subminiatus 19–19–17 keeled or smooth 144–184 56–97 1 3–4 23–26(21–24+2)
R. swinhonis 15–15–15 weakly keeled or smooth 124–165 44–74 1 2–3 19–23(17–21+2–3)
R. tigrinus 19–19–17(15) keeled or smooth 144–188 38–74 1–2 2–4 22–23(20–21+2)
Table 2 Significant cha racters of R.chiwen sp. nov. and the other known 27 species of the genus Rhabdophis.
Notes: See Material a nd methods section for the data sources.
▲: total number of maxillary teeth (the number of maxillary teeth before the diastema + the number of maxilla ry teeth behind the diastema).
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Supralabials (SL) were considered being those shields tha t were
behind the rostral and bordering the mouth gap; infrala bials
(IL) were considered being those shields tha t were behind the
mental, completely below a supralabial a nd bordering the
mouth gap. Number of supraocular (SPO), preoculars (PRO)
and postoculars (PTO) were counted as shields above, at the
anterior and posterior margin of the orbit. Loreal scales (LR)
were counted as scales between the nasal scale a nd preocula r.
Number of temporal scales (TEM) were counted as scales behind
the postoculars and between the parietal scales and suprala bials.
The sex was determined by inspection of the existence of
hemipenes.
This new species is most similar to and has long been
considered as R . pentasu pralabialis. To illuminate this problem,
22 specimens (11 males and 11 females) collected from the
type locality ( Jiulong County, JL) of R. pentasu pr alabialis were
examined and measured. Morphological characters of specimens
were analyzed using Principal Component Analysis (PCA)
(Wüster et al., 1992). Male specimens were f rom Jiulong Coun ty,
Jiguan Mounta in and Xingou Village, using the cha racters
VEN, SC, left IL, right IL, left PRO, left PTO a nd right PTO.
Female specimens were from Jiulong County and Xingou
Village, using the cha racters VEN, SC, left IL, right IL, left PTO
and right PTO. All statistical analysis were conducted with
SPSS 25.0 (IBM Inc., Armonk, U.S.A.).
3. Results
3.1. Molecular ph ylogenetic analysis We obta ined alignments
of the mitochondrial gene c yt b (1074 bp) and nuclea r gene cmos
(535 bp). Sequence data were uploaded to GenBank, availa ble
accession numbers showed in supplementa ry Ta ble S1. e best
evolution models of each partition combination are shown in
Table 3. The result show that c yt b and cmos genes consistent
phylogenetic trees were achieved by using Bayesian Inference
(BI) and Maximum likelihood RaxML (Figure 1).
The topological structures of combined c yt b and cmos
sequences phylogenetic tree are identical with the earlier study
(Takeuchi et al., 2018). Our results also strongly supported the
monophyly of Rhabdophis. The individuals from the Jigua n
Mountain and Xingou Village were placed in the genus
Rhabdophis with a high support (Figure 1). The Results also
showed that they sha re a common a ncestor with R . nuc halis, R.
leonardi, and R. pentasu pralabialis.
The specimen (HKV36838) from Hongya County, Sichua n,
China was also nested within the clade with individuals from
Jiguan Mounta in and Xingou Village.
The uncorrected p-distances between species are shown in
Table 4. The pairwise distances between species in the genus
were wide, ra nging from 0.004 to 0.198. The la rgest dispa rity
occurred between R. subminiatus from Thailand and R .
chr ysar gos from Malaysia (0.198). In comparison, the specimen of
Rhabdophis from Jiguan Mountain (CIB116092) differed from R.
pentasupralabialis from Jiulong County, Sichuan, China (GP1065,
type locality) by 0.081, and was different from R. guangdongensis
(SYSr000018) by 0.066 substitutions per site to the other species.
Moreover, the specimen (HKV36838) from Hongya County,
Sichuan, China was identified as R. nuchalis before (Alfaro and
Arnol d, 2001), but its pairwise distance with R. nuchalis (H T0854
and CIBDL1807208) were rather large, by 0.063 and 0.058
respectively.
3.2. Taxonomic conclusions e individuals from the Jiguan
Mountain and Xingou Village were placed as monophyletic
clade with strong supports (100% in PP and BP). In addition,
the p-distance between the individuals of the clade and other
species indica ted very la rge values (more tha n 0.058). Combined
with further evidence from morphology mentioned below, the
individuals from southwestern Sichuan Province represent an
undescribed species of the genus Rhabdophis. Here, we described
it as a new species.
Taxonomy
Rhabdophis chiwen sp. nov. Chen, Ding, Chen and Piao, 2019
(Figures 2, 3, 4, 6);
Rhabdophis nuchalis pentasu pralabialis: Jiang and Zhao, 1983 pp.
59–62;
Rhabdophis pentasupralabialis: Zhao, 1998, pp. 271–274; Zhao, 2006,
pp.268–269; Takeuchi et al., 2018, Figure 2 and 3, p. 10226 (part).
Diagnosis. 1) nuchal groove present, with enlarged and paired
scales on each side; 2) Dorsal body saddlebrown, DSR in 15
rows throughout, feebly keeled, the outer 1–2 rows smooth; 3)
dorsal scales typically with black margins forming some spots
and stripes, the margin of the outer row forming two faint
dorsolateral black cross-bars alongside body; 4) a black oblique
stripe below the eye, often with a black spot between the 2nd
and 3rd SL a nd a black stripe (or separated as black spots) on the
5th SL a nd the 1st TEM; 5) Eyes da rk khaki, pupils black; 6) SL
5, the 3rd and 4th touching the eye: 7) usually 7 ILs (occasionally
6 or 8), the first four contact with a nterior chin-shields; 8) 1 LR,
and TEM 1+1; 9) VEN 151-159, the outer margin of ventral scales
and several la teral rows of dorsal scales forming ventrolateral
longitudinal brownish-red coloration, with faint black spots in
Subset Best Model Site (bp) Subset Pa rtition
1 HKY+I+G 358 cyt b 1st
2 HKY+I+G 554 cmos 3rd, cyt b2nd
3 GTR+G 358 cyt b3rd
4 K80+I 392 cmos 1st,cmos2nd
Table 3 e best evolution models of each partition combina tion.
Yige PIAO et al.
A New Species of Rhabdophis
in Southwestern China
No. 2 99
Figure 1 Phylogenetic tree derived from the combined gene fragment (cmos + cyt b). Bayesian posterior proba bilities (PP) and bootstrap
probabilities (BP) from maximum likelihood a re shown at the nodes.
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ID Taxa 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
63 R. nuchalis
22 R. nuchalis 0.034
15 R. leonardi 0.045 0.047
21 R. leonardi 0.044 0.047 0.021
13 R. adleri 0.053 0.044 0.029 0.038
60 R. pentasupralabialis 0.055 0.061 0.070 0.066 0.075
16 R. pentasupralabialis 0.056 0.060 0.070 0.068 0.073 0.031
48 R. chiwen sp. nov. 0.058 0.063 0.062 0.064 0.073 0.079 0.079
69 R. chiwen sp. nov. 0.059 0.064 0.062 0.063 0.074 0.080 0.081 0.004
12 R. guangdongensis 0.047 0.053 0.057 0.055 0.058 0.074 0.071 0.065 0.066
24 R. swinhonis 0.064 0.072 0.073 0.070 0.084 0.091 0.089 0.086 0.085 0.078
40 R. swinhonis 0.076 0.082 0.086 0.084 0.096 0.097 0.090 0.086 0.086 0.089 0.066
47 R. adleri 0.132 0.126 0.131 0.132 0.131 0.134 0.128 0.130 0.132 0.117 0.131 0.136
34 R. callichromus 0.124 0.119 0.127 0.122 0.131 0.130 0.130 0.121 0.121 0.119 0.123 0.133 0.122
18 R. nigrocinctus 0.120 0.120 0.124 0.130 0.096 0.129 0.125 0.127 0.128 0.113 0.128 0.129 0.135 0.140
31 R. nigrocinctus 0.135 0.138 0.141 0.138 0.115 0.145 0.143 0.140 0.140 0.134 0.141 0.141 0.149 0.149 0.050
6R. lateralis 0.140 0.133 0.144 0.140 0.129 0.146 0.142 0.137 0.140 0.126 0.144 0.140 0.152 0.151 0.140 0.150
26 R. tigrinus 0.131 0.128 0.138 0.136 0.129 0.142 0.137 0.127 0.130 0.123 0.153 0.151 0.158 0.152 0.146 0.156 0.065
25 R. formosanus 0.135 0.134 0.140 0.138 0.129 0.141 0.140 0.141 0.139 0.128 0.141 0.148 0.168 0.160 0.149 0.161 0.090 0.088
11 R. himalayanus 0.148 0.125 0.144 0.134 0.132 0.145 0.145 0.143 0.147 0.135 0.130 0.140 0.152 0.148 0.152 0.157 0.153 0.157 0.151
29 R. subminiatus 0.152 0.147 0.150 0.153 0.150 0.156 0.156 0.159 0.162 0.149 0.158 0.156 0.169 0.170 0.166 0.177 0.172 0.165 0.170 0.155
32 R. subminiatus 0.158 0.163 0.149 0.153 0.156 0.165 0.166 0.157 0.158 0.155 0.151 0.159 0.171 0.165 0.172 0.178 0.178 0.184 0.180 0.169 0.111
30 R. chrysargos 0.190 0.187 0.184 0.184 0.177 0.189 0.188 0.181 0.182 0.179 0.179 0.178 0.194 0.197 0.191 0.197 0.188 0.182 0.196 0.176 0.197 0.198
42 R. conspicilatus 0.171 0.167 0.167 0.169 0.160 0.168 0.169 0.166 0.168 0.176 0.165 0.169 0.186 0.181 0.173 0.182 0.179 0.181 0.179 0.181 0.186 0.189 0.166
Table 4 Uncorrected p-distances between Rhabdophis species based on 1074 base pa irs from the mitochondrial genes cyt b. Specimens of R. chiwen sp. nov. are in bold font. ID = Sample ID in Ta ble
S1.
Yige PIAO et al.
A New Species of Rhabdophis
in Southwestern China
No. 2 101
the middle of each ventral scale a nd between scales which line a
black stripe; 10) anal divided; SC 45–59; 11) PRO 1 (rarely 2) and
PTO usually 3 (occasionally 2); 12) medium-sized body (SVL of
adult males: 404–431 mm and adult females: 409–476 mm); 13)
tail moderate and longer in males than in females (adult males:
10 4-131 m m, a du lt fema le s: 91–107 mm ).
Etymology. e species name of the new species “chiwen” is in
reference to the ninth son of Loong in ancient Chinese myth
who likes eating fire, and indicates the firefly-eating habit
of this new species (Yoshida et al., 2020). Its common name is
suggested as “Chiwen Keelback” in English and “ 螭吻颈槽蛇 ”
in Chinese.
Holotype. C I B11 6 09 2 , an adult male, collected from Jiguan
Mountain, Chongzhou City, Sichuan Province (30°45'57.64" N,
103°14'3.52" E, a nd 1846 m a.s.l.) collected on 1st July 2019 by Li
Ding (Figure 2 a nd 3).
Paratypes. CIB116093, an adult male, collected from the sa me
locality as holotype on 1st July 2019 by Li Ding. C I B11 6 0 9 4 -
95, two specimens, an adult male a nd an adult female, collected
from Xingou Village, Tianquan County, Sichua n Province
(29°55'42.39" N, 102°23'9.19" E, and 1461 m a.s.l.) collected on 22nd
July 2018 by Li Ding a nd Zening Chen.
3.3. Description of the holotype Adult male with TL 540
mm (SVL 422 mm a nd TaL 118 mm). TaL/TL 21.9%. Body
elongated a nd cylindrical. Nuchal groove present; seven scales
on each side of groove more or less distinctly enlarged and
paired. DSR in 15 rows throughout, feebly keeled, the outer 2
rows smooth. VEN 152; anal divided; SC 56, paired, final spine
present a t tip of tail.
Head oval and distinctly wider than neck, HL 13.67 mm,
2.53% of TL; HW 9.10 mm, HL/HW 1.50. Eyes are relatively
large, with rounded pupil, EW 2.95 mm, EW/HL 21.58%. Rostral
semi-circular when viewed from the front, wider than long,
barely visible from above. Nostrils large and open la terally,
nasal completely divided, in contact with rostral. Internasals
slightly shorter tha n prefrontals. Prefrontals wide, bending
to the loreal region. Frontal slightly pentagonal, longer than
wide, equal to the distance from the rostral to frontal. LR 1,
longitudinally longer. SPO 1; PRO 1; PTO left 2 a nd right 3;
subocular a bsent. SL 5, the 3rd and 4th touching the eye, with
a black oblique stripe below the eye, a black spot between the
2nd and 3rd SL and a black stripe on the 5th SL a nd the 1st
TEM. Mental triangular, width approximately twice of length.
IL left 7 and right 6, the first one in contact with each other
behind the mental and the first four contact with a nterior chin-
shields, the 4th to 6th infralabials contact with posterior chin-
shields. At the left side, the 7th infralabial slightly longer than
the 6th infralabial, but much narrower, width about 1/3 of its
height. TEM 2 each side, a nterior temporal single and posterior
temporal single.
Hemipenis bilobed, extends to the 11th subcaudal, forked
at the level of the 9th subcaudal. When wholly everted, the
hemipenes have small and hard spines resembling hooks
covering the entire orga n with the highest density of spines on
the tip. e length of each lobe is about 1/4 of the hemipenis.
Basal big spine absent, skin shallow cupped. The centripetal
sulcus bifurcates at the fork and extends to the tip of per lobe.
Lips evident a nd smooth (Figure 4).
Coloration of specimens in life. Dorsal body saddlebrown
with rusty red. Dorsal scales are typically with black margins
forming some sca ttered spots a nd stripes (for holotype, those
dispersed black spots forming a black stripe a round the neck,
see Figure 3 A, C, E), the margin of the outer row forming two
faint dorsolateral black cross-bars alongside body (Figure 2 A;
Figure 5 A, C). Coloration of ventral is seashell and the outer
margin of ventral scales and several lateral rows of dorsal scales
forming ventrolateral longitudinal brownish-red coloration,
with faint black spots in the middle of each ventral scale and
between scales at the anterior part of body which line a black
stripe and merge into black patches covering the whole scale
Figure 2 Dorsal (A) and ventral (B) view of the holotype of Rhab-
dophis chiwen sp. nov. (CIB116092). Photo by Shengchao Shi.
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with dispersed brownish-red spots a t the posterior part of body
(Figure 2 B). Hatchlings similar to adults except for darker
dorsal coloration, a yellow stripe present on nape, separated by
the nuchal groove (Figure 5 D); ventral scales typically shallow
black.
Coloration of specimens in preservative. Coloration pattern
in preservative similar to alive, but the faint black spots in
the middle of each ventral scale may become much dispersed,
forming large black patches alongside the ventral surface after
one year in preservative.
3.4 Variation Other specimens generally resemble the
holotype except the following characters (Ta ble 1): IL rarely 8.
PRO rarely 2 a nd PTO occasional 2. SVL ranges from 409–476
mm in females a nd 404–431 mm in males. Female specimens
tend to have a shorter ta il and fewer SC than male specimens
(f o r Ta L 91–10 7 mm vers u s 10 4 –131 mm , f or SC 4 5–52 vers u s
55–59). e ventrolateral longitudinal brownish–red coloration
may be lighter (Figure 5 A, C) and faint black spots on the
ventral scales could merge into large patches and cover the
whole scale at the posterior part of body (Figure 5 B). e dorsal
Figure 3 e holotype of Rhabdophis chiwen sp. nov. (CIB116092) in life. A–E: Different views of head. C and E: showing the same 5 su-
pralabials as R. pentasupralabialis. D: ventral view of head, displaying the representa tive 7 infralabials. F: Lateral view of trunk, demonstrat-
ing several smooth outer rows. Photo by Shengchao Shi.
Yige PIAO et al.
A New Species of Rhabdophis
in Southwestern China
No. 2 103
Figure 4 e hemipenis of the holotype of Rhabdophis chiwen sp. nov. (CIB116092). Photo by Shengchao Shi.
Figure 5 Photos of Rhabdophis chiwen sp. nov.. A and B: Ventral view of a n adult female displaying the coloration varia tions, photo by
Yige Piao. C: An adult female in life. D: A yearling in life. For C a nd D, photo by Masaya Fukuda.
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scales of some specimens (such as CIB116093) are feebly keeled
even on the second outer row, with only the outer row smooth.
3.5. Com parisons Comparative data of R. chiwen with 27
known species of the genus Rhabdo phis were obtained from
litera ture (see Ma terials and methods) a nd are shown in Ta ble 2.
Currently, 11 species are known to occur in China, and
Rhabdophis chiwen can be distinguished from these species
by following cha racters (a deta iled comparison between R.
chiwen and R. pentasupralabialis is separately listed in a latter
paragraph). R. chiwen differs from R. adleri and R. himalayanus
by DSR 15 rows throughout vs. DSR 19–19–17 in R. adleri and
R. himala yanus. R. chiwen differs from R. formosanus
and R.
lateralis by DSR 15 rows throughout vs. DSR 19–19–17(15) in
R. formosanus
and R. lateralis. The new species differs from
R. guangdongensis by 5 S L s , V EN 151–159 , S C 45–59 , dorsa l
body saddlebrown vs. 6 SLs, VEN 126, SC 39, brownish–grey
coloration in R. guangdongensis. e new species is dif ferent from
R. leonardi by DSR 15 rows throughout vs. DSR 18(17)–17–15 in
R. leonardi. e new species is different from R. nigrocinctus and
R. subminiatus by DSR 15 rows throughout vs. DSR 19–19–17 in
R. nigrocinctus and R. subminiatus. e new species differs from R.
nuchalis by 5 SLs, 7(6,8) ILs, TEM 1+1, dorsal body saddlebrown
vs. 6 SLs, 8(7) ILs, TEM 1+2, olive green colora tion with black
and magneta spots in R. nuchalis (Figure 6). R. chiwen differs from
R. swinhonis by 5 SLs, 7(6) ILs, dorsal body saddlebrown vs. 6 SLs,
8(7) ILs, medium brown coloration with several rows of black
spots in R. swinhonis.
Rhabdophis chiwen, R. pentasupr alabialis, R. guangdongensis, R.
swinhonis, R . an geli and R. nuchalis have the minimal number of
dorsal scale rows in this genus, equal to 15 rows. Nevertheless, R.
chiwen ca n be readily distinguished from the other five species
and all other species of Rhabdophis by SLs, DSR, VEN, SC,
PRO, PTO or different coloration. Beca use R. guangdongensis,
R. swinhonis and R. nuchalis are distributed in China, their
compa risons with R. chiwen are listed in the previous paragraph.
e new species differs from R. angeli b y V E N 151–159 , SC 45 –5 9 ,
dorsal body saddlebrown vs. VEN 117–126, SC 39–46, brownish
coloration with a dorsolateral series of small reddish spots in R.
angeli.
For the remaining species within the genus Rhabdophis
except R. pentasupralabialis, R. chiwen sp. nov. ca n be readily
distinguished by its dorsal scales in 15 rows throughout vs.
DS R 17–17 –15 in R. auriculata; DSR 19 at midbody in R. barbouri,
R. callichromus, R. chr ysar gos, R. conspicillatus, R. lineatus and R.
spilogaster; DSR 21 a t m id bod y in R. callistus and R. chrysar goides;
DS R 1 9–19 –15(1 7) i n R. murudensis; D S R 19–1 9–17 (15 ) in R. tigrinus.
Specimens collected from Xingou Village and Jiguan
Mountain differ from specimens of R. pentasu pralabialis by: 1)
larger size (mean TL 536.83 mm) vs. mean TL 483.64 mm in R.
pentasupralabialis (Table S2); 2) saddlebrown coloration of dorsal
body vs. dark green or olive green colora tion of dorsal body
in R. pentasupralabialis; 3) the margin of the outer row forming
two faint dorsola teral black cross-ba rs alongside body vs. a bsent
in R. pentasupralabialis; 4) the outer margins of ventral scales
and several la teral rows of dorsal scales forming ventrolateral
longitudinal brownish-red coloration, with faint black spots in
the middle of each ventral scale a nd between scales which line a
black stripe vs. a bsent in R . pentasu pralabialis (for the compa rison
of ventral coloration, see Figure S1).
For male specimens, the first two principal components
accounted for 55.900% of cumula tive coefficients. Principal
component 1 (PC1) accounted for 31.464% and principal
component 2 (PC2) for 24.436%. For female specimens, the first
two principal components accounted for 64.567%. PC1 accounted
for 39.106 % a n d PC2 f o r 25.461%. T a bl e 5 displa y s w h ich
characters are important in PC1 and PC2 for male and female
specimens.
Figure 7 shows the plots of the first two principal components
Figure 6 The lateral or ventral view of R. pentasupralabialis (A),
holotype (CIB116092) of R. chiwen sp. nov. (B) and R. nuchalis (C),
displaying the 6 ILs of R. pentasupralabialis, 7 ILs of R. chiwen sp.
nov. and 8 ILs of R. nuchalis.
Yige PIAO et al.
A New Species of Rhabdophis
in Southwestern China
No. 2 105
for males (A) and females (B). e PCA for male specimens did
not reveal much differentiation among the localities, while the
female specimens could be easily distinguished from each other.
3.6. Distribution, habitat and behavior Rhabdo phis chiwen is
currently known to be distributed in several parts of Sichuan
Province, including Xingou Village of Tia nquan County,
Jiguan Mountain of Chongzhou City a nd Hongya County.
During several field surveys in Xingou Village, individuals of
this species were commonly encountered, including hatchings,
juveniles a nd adults. At more than six different field sites with
the GPS records, we speculate that Rhabdophis chiwen lives at
the altitude ra nge of 1100-2200m, typically near fa rmla nd a nd
the source of water (Figure 8). With the stomach content by
forced regurgitation and observation of feeding behaviors in
the laboratory, it has been confirmed that this species primarily
prey on earthworms and fireflies (Yoshida et al., 2020). The
defensive behavior of this species is the typical body lift
described by Mori (Mori et al. , 2 016) .
4. Discussion
The genus Rhabdophis is widely distributed among Eurasia.
Species of this genus are known to occupy a wide range of
microhabitat a nd different altitudes (Zhao, 2006). Sichuan
Province lies in the southwestern part of China, which stretches
across Palearctic Realm and Oriental Realm, and can mainly
be divided into two pa rts: the western part is the West Sichua n
High Pla tea u and the eastern part is the Sichua n Basin (Zhao,
2002). The complex na tural environment provides varied
habitats for a large quantity of species (Wang et al. , 2010).
Currently, 5 species of Rhabdo phis are reported to inha bit
Sichuan Province, i.e. R. pentasu pralabialis, R. nuchalis, R. leonardi,
R. subminiatus and R. lateralis. ere a re doubtful records of R .
nigrocinctus, because this species typically inha bits low altitude
habitat with high humidity rather than dry-hot valleys as the
recorded locality in southwestern Sichua n (Li Ding, personal
communica tion). R. chiwen was also found in Hongya County
during several field surveys, where a specimen (HKV36838) was
considered as R. nuchalis in a previous study (Alfa ro and Arnold,
20 01). With our long-term field records since 2012, it is shown
that the distribution range of R. chiwen is partly overlapped
with that of R . pentasu pralabialis and R. nuchalis. Together with
R. leonardi, these four species all primarily feed on earthworms
and form a earthworm-eating clade while most species in this
genus prey on frogs and fish (Zhao, 2006; Yoshida et al., 2020).
Intriguingly, all these earthworm-eating species can be found
in Sichuan, and it remains to be investigated whether and how
they show some niche differentia tions.
A previous study has pointed out substantial genetic
divergence within R. nigrocinctus, R. swinhonis, R. nuchalis, and
especially R. subminiatus, and suggested the possibility for several
undescribed species (Takeuchi et al., 2018). In the study of this
genus, our field records clea rly support this possibility (Li
Component matrix
Males Females
Component Component
1 2 1 2
VEN –0.759 –0.012 0.697 0.543
SC –0.351 0.452 –0.452 0.557
left IL 0.647 0.038 0.502 –0.378
right IL 0.781 –0.069 0.487 –0.756
left PRO 0.395 0.821 - -
left PTO –0.024 –0.860 0.785 0.139
right PTO 0.563 –0.295 0.742 0.435
Table 5 Component matrix of Principal Component Analysis for
male and female specimens.
Note: See Material a nd methods section for the abbreviations.
Figure 7 e plots of the first two principal components for males (A) and females (B) of specimens.
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Ding, personal communication). It is sure that the western and
southern pa rts of Sichuan Province still provide shelters for
certa in cryptic species of Rhabdo phis.
Acknowledgements This study was supported by the
Biodiversity Survey, Observa tion and Assessment Progra mme
(2019–2023) of Ministry of Ecology and Environment of China
to Li DING and Yanqing WU; gra nts of the National Natural
Science Foundation of China (No. 31301882, No. 31970423) to
Qin Chen, a nd Science and Technology Foundation of Sichuan
(No.2018SZ0335) to Qin CHEN.; gra nts from Japa n-China Joint
Research Project (2014–2016) between the Japan Society for
the Promotion of Science ( JSPS) and National Natural Science
Foundation of China (NSFC, 31411140033) to Yezhong TANG
and Akira MORI, and grants from JSPS KAKENHI Grant
Numbers JP26440213, JP17H03719, and JP18KK0205 to Akira
MORI. In addition, we are much indebted to the Museums of
Herpetology in Chengdu Institute of Biology and their staff
for their help a nd permission to examine preserved specimens
under their care.
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How to cite this article:
Piao Y. G., Chen Z. N., Wu Y. Q., Shi S. C., Takeuchi H., Jono T., Fukuda M., Mori A., Tang Y. Z., Chen Q., Ding L.
A New Species of the Genus Rhabdophis Fitzinger, 1843 (Squama ta: Colubridae) in Southwestern Sichuan, China.
Asian Herpetol Res, 2020, 11(2): 95–107. DOI: 10.16373/j.cnki.ahr.190068
Handling Editor: Chen YANG
Appendix
Figure S1 e ventral view of R. pentasupralabialis (CIB10668) (A) from type locality and R. chiwen sp. nov. (CIBDL1807137) (B) from Xin-
gou Village, displaying the black spots and pa tches on the ventral scales in R. chiwen sp. nov. vs. absent in R. pentasupralabialis. Both speci-
mens were kept in preservative for more tha n one year.
Sample ID Species Individual No. Country and Locality cyt b cmos
1Amphiesma stolatum CAS206560 Bago Div., Myanma r AF471030 AF471097
2Natrix natrix LSUMZ41506 Sheppey Island, Kent, United Kingdom AF471059 AF471121
3Rhabdophis swinhonis KUZR18977 Taiwa n, China AB842176 AB861888
4Rhabdophis lateralis_2 GP613 China GQ281785 JQ687444
5Rhabdophis tigrinus GP1195 Huangshan, Anhui, China KF765395 KF765390
6Rhabdophis tigrinus SCUM090004 Tianshui, Ga nsu, China KF765396 KF765391
7Rhabdophis tigrinus SCUM090001 Longnan, Ga nsu, China KF765397 KF765392
8Rhabdophis subminiatus RDQ200905366 Baoshan, Yunnan, China KF765398 KF765393
9Rhabdophis subminiatus SCUM090014 Panzhihua, Sichuan, China KF800927 KF765394
10 Rhabdophis subminiatus SICAU090029 Hong Kong, China KF800928 KF800918
11 Rhabdophis himalayanus CAS224420 Kachin State, Mya nmar KF800929 KF800919
12 Rhabdophis guangdongensis SYSr000018 Aizhai, Renhuay, Guangdong, China KF800930 KF800920
13 Rhabdophis adleri WJC20090801 Ha inan, China KF800931 KF800921
14 Rhabdophis leonardi RDQ200905367 Dali, Yunnan, China KF800932 KF800922
15 Rhabdophis leonardi SCUM090009 Panzhihua, Sichuan, China KF800933 KF800923
16 Rhabdophis pentasupralabialis GP1065 Jiulong, Sichuan, China KF800934 KF800924
17 Rhabdophis nuchalis SICAU090001 Shennongjia, Hubei, China KF800935 KF800925
18 Rhabdophis nigrocinctus CAS215280 Shan State, Myanmar KF800936 KF800926
19 Rhabdophis nigrocinctus_3 HT0845 China LC325298
20 Rhabdophis himalayanus_1 HT0847 China LC325299 LC325746
21 Rhabdophis leonardi_1 HT0851 China LC325300 LC325747
22 Rhabdophis nuchalis_4 HT0854 China LC325301 LC325748
23 Rhabdophis lateralis_1 HT0855 China LC325302
24 Rhabdophis swinhonis_1 HT0021 Ta iwan, China LC325303 LC325749
25 Rhabdophis formosanus_1 HT0033 Ta iwan, China LC325304 LC325750
26 Rhabdophis tigrinus_1 HT0098 Japan LC325305 LC325751
27 Rhabdophis tigrinus_2 HT0177 Japan LC325306 LC325752
28 Rhabdophis nigrocinctus_1 HT0253 ailand LC325307 LC325753
29 Rhabdophis subminiatus_1 HT0267 Laos LC325308 LC325754
30 Rhabdophis chrysargos HT0342 Malaysia LC325313 LC325759
31 Rhabdophis nigrocinctus_2 HT0343 ailand LC325314 LC325760
32 Rhabdophis subminiatus_2 HT0344 ailand LC325315 LC325761
33 Rhabdophis subminiatus_3 HT0345 ailand LC325316 LC325762
34 Rhabdophis callichromus_1 HT0654 Vietnam LC325324 LC325770
35 Rhabdophis callichromus_2 HT0674 Vietnam LC325325 LC325771
36 Rhabdophis subminiatus_4 HT0680 Vietnam LC325328 LC325774
37 Rhabdophis pentasupralabialis_1 HT0699 China LC325331 LC325777
38 Rhabdophis pentasupralabialis_2 HT0700 China LC325332 LC325778
39 Rhabdophis nuchalis_1 HT0701 China LC325333 LC325779
40 Rhabdophis swinhonis_2 HT0717 Ta iwan,China LC325334 LC325780
41 Rhabdophis murudensis HT0788 Malaysia LC325341 LC325787
42 Rhabdophis conspicilatus HT0791 Malaysia LC325342 LC325788
43 Rhabdophis nuchalis_2 HT0803 China LC325352 LC325798
44 Rhabdophis nuchalis_3 HT0807 China LC325353 LC325799
45 Rhabdophis pentasupralabialis_3 HT0808 China LC325354 LC325800
46 Rhabdophis adleri_1 HT0831 China LC325356 LC325802
47 Rhabdophis adleri_2 HT0832 China LC325357 LC325803
48 Rhabdophis nuchalis HKV36838 Hongya County, Sichuan, China AF402907
49 Rhabdophis chiwen sp. nov. CIB DL2014155 Jigua n Mountain, Sichuan, China MN656319 MN656340
Table S1 Information of all sequence da ta used in the study.
Sample ID Species Individual No. Country and Locality cyt b cmos
50 Rhabdophis chiwen sp. nov. CIB DL1807203 Jigua n Mountain, Sichuan, China MN656320 MN656341
51 Rhabdophis chiwen sp. nov. CIB DL1807204 Jigua n Mountain, Sichuan, China MN656321 MN656342
52 Rhabdophis chiwen sp. nov. CIB DL1807206 Jigua n Mountain, Sichuan, China MN656322 MN656343
53 Rhabdophis nuchalis CIB DLHK1 Chongqing, China MN656323 MN656344
54 Rhabdophis chiwen sp. nov. CIB DL2014M1 Jiguan Mounta in, Sichuan, China MN656324 MN656345
55 Rhabdophis chiwen sp. nov. CIB DLM415 Jigua n Mountain, Sichuan, China MN656325 MN656346
56 Rhabdophis nuchalis CIB DL2014M5 Longnan, Ga nsu, China MN656326 MN656347
57 Rhabdophis leonardi CIB DL1806011 Panzhihua, Sichuan, China MN656327 MN656348
58 Rhabdophis leonardi CIB DL1807031 Panzhihua, Sichuan, China MN656328 MN656349
59 Rhabdophis pentasupralabialis CIB DL1807061 Pa nzhihua, Sichuan, China MN656329 MN656350
60 Rhabdophis pentasupralabialis CIB DL1809267 Pa nzhihua, Sichuan, China MN656330 MN656351
61 Rhabdophis nuchalis CIB DL1807207 Ta ngjiahe, Sichuan, China MN656333 MN656354
62 Rhabdophis nuchalis CIB DL1807208 Ta ngjiahe, Sichuan, China MN656334 MN656355
63 Rhabdophis nuchalis CIB DL1807209 Ta ngjiahe, Sichuan, China MN656335 MN656356
64 Rhabdophis chiwen sp. nov. CIB116094 Xingou Village, Sichua n, China MN656336 MN656357
65 Rhabdophis chiwen sp. nov. CIB DL1804151 Xingou Village, Sichua n, China MN656337 MN656358
66 Rhabdophis chiwen sp. nov. CIB116095 Xingou Village, Sichua n, China MN656338 MN656359
67 Rhabdophis leonardi CIB DL1809265 Yingjiang, Yunnan, China MN656318 MN656339
68 Rhabdophis chiwen sp. nov. CIB116092 Jigua n Mountain, Sichuan, China MN656331 MN656352
69 Rhabdophis chiwen sp. nov. CIB116093 Jigua n Mountain, Sichuan, China MN656332 MN656353
(Continued Table S1)
Locality Number Sex SVL TL Ta L ED HL HW Ven Sc ASR MSR DSR SL-left S-right IL-left IL-right PRO PTO LR TEM
Xingou Village CIBDL1807135 Female 476 583 107 2.97 14.55 11.25 156 46 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Xingou Village CIBDL1807132 Male 427 555 128 3.05 13.39 9.22 154 58 15 15 15 2-2-1 2-2-1 8(4) 7(4) 1 3 1 2
Xingou Village CIB116094 Male 411 524 11 3 2.95 13.06 9.02 154 55 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Xingou Village CIBDL1807133 Male 409 537 128 2.63 12.95 7.95 157 58 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Xingou Village CIBDL1807138 Male 427 549 122 2.63 13.90 8.45 152 59 15 15 15 2-2-1 2-2-1 7(4) 7(4) L:2 R:1 L:1 R:3 1 2
Xingou Village CIBDL18071311 Female 422 513 91 2.86 12.60 8.84 158 47 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Xingou Village CIBDL1807134 Female 4 11 509 98 2.55 11.91 7.33 159 49 15 15 15 2-2-1 2-2-1 7(4) 6(4) 1 L:3 R:2 1 2
Xingou Village CIBDL1807139 Male 404 508 104 2.32 13.61 9.02 151 55 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Xingou Village CIB116095 Female 461 557 96 2.70 13.90 8.96 157 45 15 15 15 2-2-1 2-2-1 6(4) 7(4) 1 L:3 R:2 1 2
Xingou Village CIBDL1807137 Female 409 505 96 2.90 12.13 7.61 159 52 15 15 15 2-2-1 2-2-1 7(4) 6(4) 1 3 1 2
Jiguan Mountain CIB116092 Male 422 540 118 2.95 13.67 9.10 152 56 15 15 15 2-2-1 2-2-1 7(4) 6(4) 1 L:2 R:3 1 2
Jiguan Mountain CIB116093 Male 431 562 131 2.88 13.55 9.28 155 57 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 3 1 2
Jiulong County CIB10657 Male 382 486 104 2.17 12.08 7.96 158 57 14 15 15 2-2-1 2-2-1 5(4) 6(4) 1 3 1 2
Jiulong County CIB10660 Female 383 474 91 2.60 13.01 8.12 150 49 15 15 15 2-2-1 2-2-1 6(4) 5(4) 1 L:3 R:2 1 2
Jiulong County CIB10689 Male 360 454 94 2.38 11.77 8.51 154 56 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 L:2 R:3 1 2
Jiulong County CIB10681 Female 419 528 109 2.25 13.60 8.43 160 52 15 15 15 2-2-1 2-2-1 5(4) 5(4) 1 3 1 2
Jiulong County CIB10655 Male 370 475 105 2.25 11.38 8.06 157 59 15 15 15 2-2-1 2-2-1 7(4) 5(3) 1 L:3 R:2 1 2
Jiulong County CIB10690 Female 427 535 108 2.37 13.25 8.73 155 55 15 15 15 2-2-1 2-2-1 6(4) 5(4) 1 2 1 2
Jiulong County CIB10636 Male 367 466 99 2.17 12.88 8,71 156 55 14 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10650 Female 391 498 107 2.17 12.70 9.30 154 51 14 15 15 2-2-1 2-2-1 6(4) 5(4) 1 2 1 2
Jiulong County CIB10696 Male 339 425 86 2.03 11.44 7.90 153 55 15 15 15 2-2-1 2-2-1 6(4) 7(4) 1 3 1 2
Jiulong County CIB10654 Male 348 444 96 2.23 11.83 7.98 152 56 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10684 Female 398 500 102 2.22 12.76 8.82 152 52 14 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10823 Female 347 452 105 1.96 11.20 7.83 157 62 15 15 15 2-2-1 2-2-1 7(4) 6(4) 1 L:2 R:3 1 2
Jiulong County CIB10850 Female 390 496 106 2.33 12.95 9.15 153 57 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10800 Female 394 515 121 2.22 13.04 8.69 157 61 15 15 15 2-2-1 2-2-1 7(4) 6(4) 1 2 1 2
Jiulong County CIB10812 Male 389 501 112 2.17 12.54 8.08 159 58 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10691 Female 382 473 91 2.32 12.99 9.34 152 46 15 15 15 2-2-1 2-2-1 7(4) 8(5) 1 2 1 2
Jiulong County CIB10668 Female 418 530 112 2.09 13.67 9.21 153 53 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10652 Female 437 548 111 2.16 14.16 9.88 154 51 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10656 Male 366 467 101 2.29 12.05 7.93 155 58 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 2 1 2
Jiulong County CIB10659 Male 382 488 106 2.37 12.75 8.39 160 58 15 15 15 2-2-1 2-2-1 6(4) 6(4) 1 2 1 2
Jiulong County CIB10836 Male 338 432 94 2.21 12.23 7.68 153 60 15 15 15 2-2-1 2-2-1 6(4) 5(4) 1 3 1 2
Jiulong County CIB10841 Male 347 453 106 1.82 11.85 7.52 152 55 15 15 15 2-2-1 2-2-1 7(4) 7(4) 1 2 1 2
Table S2 Morphological characters of specimens used in the study. See Ma terial and methods section for the abbrevia tions. For the IL list, the numbers in brackets indicate the number of
infralabials touching the a nterior chin-shields.
