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A new species of Gonyosoma Wagler, 1828 (Serpentes, Colubridae), previously confused with G. prasinum (Blyth, 1854)

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A new species of the genus Gonyosoma Wagler is described from Yunnan Province, China. The new species closely resembles G. prasinum (Blyth), but it is differentiated from the latter species by the following characters: precloacal plate divided, iris blue and inside of mouth greyish-white in life. Based on phylogenetic analyses of mitochondrial cytochrome b sequence data, the new species is recovered as the sister species to G. prasinum by Bayesian Inference and Maximum Likelihood analyses. The uncorrected pairwise distance between the new species and other species of the genus Gonyosoma ranged from 11.78% to 17.07% calculated using the mitochondrial cytochrome b sequence. This discovery increases the number of Gonyosoma species to seven.
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A new species of Gonyosoma Wagler, 1828 (Serpentes, Colubridae),
previously confused with G. prasinum (Blyth, 1854)
Shuo Liu1, Mian Hou2, Ye Htet Lwin3, Qiaoyan Wang4, Dingqi Rao5
1 Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, 650223, China
2 College of Continuing (Online) Education, Sichuan Normal University, Chengdu, 610066, China
3 Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, Nay Pyi Taw, 05282, Myanmar
4 Research Institute of Xishuangbanna National Nature Reserve, Jinghong, 666100, China
5 Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, 650201, China
http://zoobank.org/03620F93-FD7A-4396-B158-554FD9866350
Corresponding author: Dingqi Rao (raodq@mail.kiz.ac.cn)
Academic editor: Alexander Haas
Received
27 March 2021
Accepted
31 May 2021
Published
22 June 2021
Abstract
A new species of the genus Gonyosoma Wagler is described from Yunnan Province, China. The new species closely resembles
G. prasinum (Blyth), but it is dierentiated from the latter species by the following characters: precloacal plate divided, iris blue and
inside of mouth greyish-white in life. Based on phylogenetic analyses of mitochondrial cytochrome b sequence data, the new species
is recovered as the sister species to G. prasinum by Bayesian Inference and Maximum Likelihood analyses. The uncorrected pairwise
distance between the new species and other species of the genus Gonyosoma ranged from 11.78% to 17.07% calculated using the
mitochondrial cytochrome b sequence. This discovery increases the number of Gonyosoma species to seven.
Key Words
Htamanthi, morphology, phylogeny, systematics, taxonomy, Yunnan
Introduction
Once considered members of a single genus, Elaphe
Fitzinger, the ratsnakes have been divided into a number
of genera including Gonyosoma (Burbrink and Lawson
2007; Chen et al. 2017). The genus Gonyosoma current-
ly includes six recognized species, namely G. boulengeri
(Mocquard, 1897); G. frenatum (Gray, 1853); G. jansenii
(Bleeker, 1859); G. margaritatum Peters, 1871; G. oxy-
cephalum (Boie, 1827) and G. prasinum (Chen et al.
2014, 2017; Uetz et al. 2021).
Gonyosoma prasinum, a wholly green, arboreal snake
(Schulz 1991, 1996; Das 2002, 2012), its type locality
is in Assam, Northeastern India (Blyth 1854; Boulenger
1894; Smith 1943), and has been considered to be widely
distributed in India, Bhutan, Myanmar, Thailand, Malay-
sia, Laos, Vietnam, and China (Schulz 1996; Zhao et al.
1998; Wu et al. 2002; Vogel and Pauwels 2004; Pauwels
et al. 2006; Zhao 2006; Ziegler et al. 2007; Yang and
Rao 2008; Hecht et al. 2013; Pham et al. 2014; Luu et al.
2020; Nguyen et al. 2020; Wang et al. 2020; Wangyal et
al. 2020; Uetz et al. 2021).
During our eldworks in southern China and northern
Myanmar from 2019 to 2020, some specimens of Gon-
yosoma cf. prasinum were collected, six from Yunnan
Province, China, and two from Htamanthi Wildlife Sanc-
tuary, northwestern Myanmar, relatively close to the type
locality of G. prasinum. The specimens from China and
Myanmar showed some dierences in scale counts and
coloration in life. In addition, phylogenetically, the se-
quences of the specimens from China and from Myanmar
were placed in two separate clades and had an obvious
genetic divergence. Therefore, we treated the specimens
from China and from Myanmar to be two distinct species.
The question was, which one is the true Gonyosoma
prasinum? As no molecular data for the type specimens
Evolutionary Systematics. 5 2021, 129–139 | DOI 10.3897/evolsyst.5.66574
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Shuo Liu et al.: A new snake species from Chian130
is available, we can only solve this problem through mor-
phological characteristics. However, we cannot judge
this problem from the original description (Blyth 1854)
of G. prasinum as it is too short and not detailed enough.
Since most of the subsequent descriptions and photo-
graphs (e.g., Boulenger 1894; Smith 1943; Schulz 1996;
Cox et al. 1998; Zhao et al. 1998; Zhao 2006; Yang and
Rao 2008; Das 2010, 2012; Shi et al. 2011; Purkayastha
2013; Qi 2019) of G. prasinum are either quoted from
previous publications or based on specimens whose
collection sites are distant from the type locality, these
specimens may not be the true G. prasinum, so these data
cannot be used here. We can only identify this species
according to the descriptions and photographs (e.g., Das
2013; Khandekar et al. 2021) of specimens originating
from its type locality and its vicinities.
Materials and methods
Sampling
Field surveys in Yunnan, China, were conducted under
the permits of Xishuangbanna National Natural Re-
serve Management Bureau and Wuliangshang Nation-
al Nature Reserve Management Bureau. Field survey
in Northern Myanmar was undertaken at the invitation
of the Republic of the Union of Myanmar, Ministry of
Natural Resources and Environmental Conservation,
Forest Department, Forest Research Institute. Four spec-
imens (KIZ2019025–KIZ2019028) were collected from
Mengla County, Yunnan Province, China, in April to
May 2019, one specimen (KIZ20200729) was collect-
ed from Zhenyuan County, Yunnan Province, China, in
July 2020, one specimen (KIZ20200903) was collected
from Menglian County, Yunnan Province, China, in Sep-
tember 2020, and two specimens (SEABRI2019120043,
SEABRI2019120075) were collected from Htamanthi
wildlife sanctuary, Sagaing, Myanmar, in December
2019. Specimens were collected by hand, and photo-
graphs were taken to document color pattern in life prior
to euthanasia. Liver tissues were stored in 99% ethanol
and snakes were preserved in 75% ethanol. The speci-
mens from China were deposited in Kunming Institute
of Zoology, Chinese Academy of Sciences (KIZ) and the
specimens from Myanmar were deposited in Southeast
Asia Biodiversity Research Institute, Chinese Academy
of Sciences (SEABRI).
Molecular data and phylogenetic analyses
All species of the genus Gonyosoma were included in
the study. Homologous sequences were obtained from
GenBank. Six samples from China and two sample from
Myanmar were incorporated in the analysis, and the new
sequences have been deposited in GenBank. Elaphe tae-
niura (Cope) and Coelognathus radiatus (Boie) were
used as outgroups. All the GenBank accession numbers
for taxa used in this study were listed in Table 1.
DNA was extracted using DNeasy Blood and Tissue
kit (Qiagen, Germany) following the manufacturer’s in-
structions. A fragment of mitochondrial cytochrome b
(cytb) gene was amplied using the primer pair L14910:
5’–GACCTGTGATMTGAAAACCAYCGTT–3’
and H16064: 5’–CTTTGGTTTACAAGAACAAT-
GCTTTA–3’ (Burbrink et al. 2000). PCR amplications
were performed in 25 μl reactions by using the follow-
ing cycling conditions: initial denaturation for 120 s at
95 °C, followed by 35 cycles: denaturation at 94 °C for
40 s, annealing at 48.5 °C for 25 s, elongation at 72 °C for
15 s, and then nalized with elongation step of 120 s at
72 °C, with the PTC-100 thermal cycler (BioRad, USA).
The products were puried by using the DNA Agarose
Gel Extraction Kit (Omega, USA) according to the man-
ufacturer’s instructions. Puried PCR products were se-
quenced using the same PCR primers. Sequencing was
completed by Beijing Qingke New Industry Biotechnol-
ogy Co., Ltd. Sequences were edited and manually man-
aged using SeqMan in Lasergene 7.1 (DNASTAR Inc.,
Madison, WI, USA) and MEGA X (Kumar et al. 2018).
Sequences were aligned using ClustalX 2.0 (Thomp-
son et al. 1997) with the default parameters. Pairwise
distances between species were calculated in MEGA
X (Kumar et al. 2018). The best substitution models
HKY+F+I+G4 and GTR+F+I+G4 were selected for
Bayesian inference and Maximum Likelihood analysis,
respectively, using the Akaike Information Criterion
(AIC) in ModelFinder (Kalyaanamoorthy et al. 2017).
Table 1. Sequences of cytb gene used in this study.
Species Voucher Locality Accession
Coelognathus
radiatus
CHS556 Wenshan, Yunnan, China MK201411
Elaphe taeniura CHS203 Xiangcheng, Sichuan,
China
MK201333
Gonyosoma
boulengeri
YPX11032 /AF471053
CHS242 Mengzi, Yunnan, China MK201360
CHS243 Hainan, China MK201361
Gonyosoma
frenatum
YPX11027 /KF669250
HS11038 /KF669251
EXTRACTION /KF669246
/ / DQ902110
CHS138 Huangshan, Anhui, China MK201289
CHS139 Huangshan, Anhui, China MK201290
Gonyosoma
jansenii
/Sulawesi DQ902113
KU 321724 Zamboanga, Mindanao,
Philippines
KC010343
Gonyosoma
margaritatum
/ / KM870886
Gonyosoma
oxycephalum
/ / AF471084
ROM 37622 /KX694870
Gonyosoma
prasinum
SEABRI2019120043 Htamanthi, Sagaing,
Myanmar
MZ322864
SEABRI2019120075 MZ322863
Gonyosoma
coeruleum sp. nov.
KIZ2019025 Mengla, Yunnan, China MZ322870
KIZ2019026 MZ322869
KIZ2019027 MZ322868
KIZ2019028 MZ322867
KIZ20200729 Zhenyuan, Yunnan, China MZ322866
KIZ20200904 Menglian, Yunnan, China MZ322865
Evolutionary Systematics 5 2021, 129–139
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131
Bayesian inference was performed in MrBayes 3.2.7
(Ronquist et al. 2012). Two runs were performed simul-
taneously with four Markov chains starting from random
tree. The chains were run for 10,000,000 generations and
sampled every 1000 generations. The rst 25% of the
sampled trees was discarded as burn-in after the stan-
dard deviation of split frequencies of the two runs was
less than a value of 0.01, and then the remaining trees
were used to create a 50% majority-rule consensus tree
and to estimate Bayesian posterior probabilities. Maxi-
mum Likelihood analysis was performed in RaxmlGUI
2.0 (Silvestro and Michalak 2012), nodal support was
estimated by 1,000 rapid bootstrap replicates.
Morphology
Body and tail lengths were measured with a ruler to the
nearest 1 mm (Jablonski et al. 2019). Scale counts were
taken following Janssen et al. (2019). Ventral scales
were counted according to Dowling (1951). Bilateral
scale counts were given as left / right order. Abbrevia-
tions of morphological characters are as follows: Atem,
number of anterior temporals; DSR, dorsal scale rows
number at one head length posterior to the head – num-
ber of dorsal scale rows at midbody – number of dorsal
scale rows at one head length anterior to the vent, re-
spectively; IL, infralabials (counted on lower lips); Lor,
loreals; Lor / eye, loreal scale touching the eye (yes or
no); PostOc, postoculars; Prec, precloacal plate (corre-
sponds to term anal plate in older literature) (single or
divided); PreOc, preoculars; PTem, number of posterior
temporals; SL, supralabials (counted on upper lips); SL /
orbit, number of supralabials entering orbit; SubC, num-
ber of subcaudal scales; SVL, snout-vent length (from
tip of snout to vent); TaL, tail length; TaL / TL, ratio of
tail length / total length; TL, total length; Ven, number
of ventral scales.
Results
The topologies derived from Bayesian inference and
Maximum Likelihood analysis were consistent (Fig. 1).
The sequences of the specimens from China formed a
separate clade sister to the sequences of the specimens
from Myanmar, however, the nodal supports were low.
The average uncorrected pairwise distance (p-distance)
between the sequences of the specimens from China
and the sequences of the specimens from Myanmar is
11.78%. (Table 2).
Morphological measurements, scale counts and color-
ations in life were presented in Table 3. The coloration of
the iris of the specimens from Myanmar and from China
in life are obviously dierent, the coloration of the iris of
the specimens from Myanmar in life is greenish yellow
(Figs 4B, 5C, D) while the colorations of the iris of the
specimens from China are blue (Figs 4A, 5A, B). Besides
this dierence, the precloacal plates of the specimens
from Myanmar are single (Fig. 4F) while the precloacal
plates of the specimens from China are divided (Fig. 4E).
According to the photographs (Khandekar et al. 2021) of
Gonyosoma prasinum in life from the type locality and
the vicinities of the type locality, the colorations of the
iris are all greenish-yellow. In addition, according to the
description of Gonyosoma prasinum based on a specimen
from Namchik, Arunachal Pradesh (Das 2013), which is
close to the type locality, the precloacal plate is single.
Therefore, we consider the specimens from Myanmar to
be conspecic with the true G. prasinum and the speci-
mens from China to belong to a new species.
Gonyosoma coeruleum sp. nov.
http://zoobank.org/81045CF9-665A-4E9C-A563-4794208D0A40
Figs 2, 3, 4A, C, E, 5A, B, 7B
Holotype. KIZ2019028, adult female, Mengla County,
Xishuangbanna Autonomous Prefecture, Yunnan Prov-
ince, China, 29 April 2019, 21°32'12"N, 101°32'51"E,
900 m a.s.l.
Paratypes. KIZ2019025, one adult female and
KIZ2019026–KIZ2019027, two adult males, Mengla
County, Xishuangbanna Autonomous Prefecture, Yunnan
Province, China, 4 May 2019, 21°55'9"N, 101°32'12"E,
890 m, a.s.l. KIZ20200729, adult female, collected from
Zhenyuan County, Puer City, Yunnan Province, China,
29 July 2020, 24°3'37"N, 101°3'43"E, 1,240 m a.s.l.
KIZ20200904, one juvenile, collected from Menglian
County, Puer City, Yunnan Province, China, 3 September
2020, 22°10'16"N, 99°18'31"E,1,200 m a.s.l.
Etymology. The specic epithet “coeruleum” is the
neutral gender of the Latin adjective coeruleus (a, um)
meaning “blue”, and is given in reference to the color-
ation of the iris of this species.
Diagnosis. Body size medium (SVL 656–833mm in
adults); body slender, head elongated and distinct from
neck; large eyes with round pupil; tail long (23–28% of
total length) and slender; dorsal scales in 19-19-15 rows,
7–11 rows of mid-dorsal scales keeled; single preocular;
Table 2. Average uncorrected p-distances (%) between mem-
bers of Gonyosoma and outgroups calculated from cytb gene
sequences.
1 2 3 4 5 6 7 8
1 Gonyosoma coeruleum
sp. nov.
2 Gonyosoma
boulengeri
13.01
3 Gonyosoma frenatum 13.35 7.56
4 Gonyosoma jansenii 17.07 15.63 16.88
5 Gonyosoma
margaritatum
12.33 12.51 12.41 16.91
6 Gonyosoma
oxycephalum
15.77 15.32 16.36 7.75 15.98
7 Gonyosoma prasinum 11.78 12.89 12.65 17.69 12.71 15.96
8 Coelognathus radiatus 19.32 18.40 18.15 19.49 18.97 19.66 17.35
9 Elaphe taeniura 16.27 15.56 15.59 16.21 16.07 16.92 16.32 15.90
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Shuo Liu et al.: A new snake species from Chian132
Figure 1. The consistent phylogram inferred from Bayesian Inference and Maximum Likelihood analyses based on cytb gene se-
quences. Numbers before slashes indicate Bayesian posterior probabilities (values below 0.9 were not shown) and numbers after
slashes indicate bootstrap support for Maximum Likelihood analyses (values below 60 were not shown).
Table 3. Measurements (in mm), scalation data, and coloration of Gonyosoma coeruleum sp. nov. and G. prasinum. For abbrevia-
tions see Materials and methods.
Gonyosoma coeruleum sp. nov. Gonyosom prasinum
KIZ2019028
Holotype
KIZ2019025
Paratype
KIZ2019026
Paratype
KIZ2019027
Paratype
KIZ20200729
Paratype
KIZ20200904
Paratype
SEABRI2019120043 SEABRI2019120075
Sex Female Female Male Male Female Juvenile Male Male
SVL 814 656 833 719 696 388 487 577
TL 1058 863 974+ 992 918 511 656 779
TaL 245 207 141+ 273 222 123 169 202
TaL / TL 0.23 0.24 0.14+ 0.28 0.24 0.24 0.26 0.26
SL 8/9 9/9 9/9 9/9 9/9 9/9 9/9 9/9
SL / orbit 3-5/4-6 4-6 4–6 4–6 4–6 4–6 4–6 4–6
IL 10/10 10/10 10/10 10/10 11/10 10/10 11/10 10/10
PreOc 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1
PostOc 2/2 2/2 2/2 2/2 2/2 2/2 2/2 2/2
Lor 1/1 1/1 1/1 1/1 1/1 1/1 1/1 1/1
Lor / eye no No no no no no no no
Atem 2/2 2/2 2/2 1/1 1/2 1/1 1/1 1/2
PTem 2/2 3/2 2/2 2/2 2/2 2/2 1/1 2/2
DSR 19-19-15 19-19-15 19-19-15 19-19-15 19-19-15 19-19-15 19-19-13 19-19-15
Ven 202 199 196 190 189 195 194 193
SubC 89 103 44+ 106 93 105 104 105
Prec Divided Divided Divided Divided Divided Divided Single Single
Body color Green Green Green Green Green Green Green Green
Tongue color Brownish
yellow
Brownish
yellow
Brownish
yellow
Brownish
yellow
Brownish
yellow
Brownish
yellow
Brownish yellow Brownish yellow
Inside of mouth Greyish white Greyish white Greyish white Greyish white Greyish white Greyish white Pink Pink
Iris color Blue Blue Blue Blue Blue Blue Greenish yellow Greenish yellow
Evolutionary Systematics 5 2021, 129–139
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Figure 2. Type series of Gonyosoma coeruleum sp. nov. in preservative. A dorsal view B ventral view.
two postoculars; one or two anterior temporals and two
or three posterior temporals; 189–202 ventral scales; 89–
106 paired subcaudals; precloacal plate divided. Dorsal
surface bright green with brownish-yellow tip of tail, iris
blue, inside of mouth greyish white; tongue brownish yel-
low with black tips.
Description of the holotype. Head elongate, distinct
from the neck, attened, longer than wide, narrowed
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Shuo Liu et al.: A new snake species from Chian134
Figure 3. Head views of the holotype (KIZ2019028) of Gonyosoma coeruleum sp. nov. in preservative. A dorsal B ventral C right
D left.
anteriorly; nostril lateral; eye large, pupils round; ros-
tral triangular, broader than high, visible from above;
nasal divided into two scales; two internasals, wider
than high, bordered by two large prefrontals posteri-
orly; frontal single, enlarged, pentagonal, narrowed
posteriorly; parietals longer than wide, in contact with
each other; supralabials 8/9, rst and second in contact
with the prenasal and postnasal, third and fourth enter-
ing orbit on left side, fourth and sixth entering orbit on
right side, eighth largest; infralabials 10/10, rst pair
in broad contact with each other, rst to fth in contact
with anterior pair of chin shields; anterior and posteri-
or pairs of chin shields elongate, second pair meeting
in midline; preocular 1/1; postoculars 2/2, lower ones
smaller, bordering anterior temporals; anterior tempo-
rals 2/2, posterior temporals 2/2. SVL 814 mm; TaL 245
mm; TaL/TL 0.23; DSR 19-19-15, nine rows keeled in
the vertebral region, otherwise smooth; ventrals 202
with a lateral keel; subcaudals 89, paired; precloacal
plate divided.
Color of holotype in life. Dorsal surface bright green
with brownish-yellow tip of tail, reticulate pattern con-
sisting of yellow, black, and white on the interstitial
skin; upper lips yellowish-green; anterior ventral surface
greenish-white and posterior ventral surface light green;
tip of tail brownish-yellow on its ventral surface; ventrals
outside the lateral keel usually yellowish-white; iris blue;
pale grey inside of mouth; tongue brownish-yellow with
black tips.
Variations. The paratypes resemble the holotype in
most aspects except that the rows of mid-dorsal scales
keeled vary from seven to eleven, anterior temporals
vary from one to two, and posterior temporals vary
from two to three in paratypes; moreover, the male
paratype KIZ2019027 has a relatively longer tail (TaL/
TL 0.28).
Distribution. Gonyosoma coeruleum sp. nov. is cur-
rently known from Xishuangbanna Autonomous Pre-
fecture and Puer City, Yunnan Province, China (Fig. 6),
it is probably also distributed in other parts of Yunnan
Province and Sichuan, Guizhou, and Hainan Province,
China, as well as Southern Myanmar, Thailand, Laos,
and Vietnam.
Natural history. The specimens from Mengla County
were collected on big trees bordering rivers, the specimen
from Zhenyuan County was collected on a big tree on
the side of a small road in forest, and the specimen from
Menglian County was collected on a small tree bordering
Evolutionary Systematics 5 2021, 129–139
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135
Figure 4. Comparisons of the colors of iris, the colors of inside of mouth, and the precloacal plates. A, C, and E the holotype
(KIZ2019028) of Gonyosoma coeruleum sp. nov. B, D, and F the specimen (SEABRI2019120043) of G. prasinum.
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Shuo Liu et al.: A new snake species from Chian136
Figure 5. The specimens of Gonyosoma coeruleum sp. nov. and G. prasinum in life. A the female paratype (KIZ20200729)
of Gonyosoma coeruleum sp. nov. B the juvenile paratype (KIZ20200904) of Gonyosoma coeruleum sp. nov. C the specimen
(SEABRI2019120043) of G. prasinum from Myanmar D the specimen (SEABRI2019120075) of G. prasinum from Myanmar.
a stream. All specimens were found at night while they
were asleep on tree branches, what shows that this spe-
cies is diurnal. Through direct observation, we found that
they like to feed on small rodents, and whether they also
prey on other animals is unknown.
Comparisons. Gonyosoma coeruleum sp. nov. can
be distinguished from G. boulengeri in lacking a na-
sal appendage (vs. rostral distinct from the nasal ap-
pendage), and no dark stripe behind the eye (vs. an in-
distinct dark stripe behind the eye) (Mocquard 1897;
Smith 1943).
Gonyosoma coeruleum sp. nov. is distinguishable from
G. frenatum based on its single, distinct loreal (vs. lore-
al united with the prefrontal), and no black streak along
the side of the head (vs. a black streak along the side of
the head above the supralabials) (Gray 1853; Boulenger
1894; Smith 1943).
Gonyosoma coeruleum sp. nov. can be separated from
G. jansenii by having 19 midbody dorsal scale rows (vs.
23–25), and dorsal surface uniform bright green with
brownish-yellow tip of tail (vs. olive or yellowish-brown,
entirely black posteriorly and on the tail) (Bleeker 1859;
Boulenger 1894).
From Gonyosoma margaritatum, Gonyosoma coeru-
leum sp. nov. can be dierentiated by its dorsal surface
uniform bright green with brownish-yellow tip of tail
(vs. black, each scale with a yellowish green spot, or
green with black borders to the scales, hinder part of
body and tail with bright orange rings), and no black
streak on each side of the head (vs. a black streak on
each side of the head behind the eye) (Peters 1871; Bou-
lenger 1894).
From Gonyosoma oxycephalum, Gonyosoma coeru-
leum sp. nov. can be dierentiated by the dierent colors
of tail (green with brownish-yellow tip vs. the whole tail
light chestnut or bu-red or yellowish brown), no blackish
stripe along the side of the head (vs. an indistinct blackish
stripe along the side of the head immediately above the
supralabials), and the number of midbody dorsal scale,
19 rows (vs. 23–27 rows) (Boie 1827; Boulenger 1894;
Smith 1943).
Gonyosoma coeruleum sp. nov. closely resembles G.
prasinum, but the colorations of their iris in life are obvi-
ously dierent, the iris of Gonyosoma coeruleum sp. nov.
is blue (Fig. 4A) while the iris of G. prasinum is green-
ish-yellow (Fig. 4B; Khandekar et al. 2021). The colours
of the inside of mouth are slightly dierent, the inside
of mouth of Gonyosoma coeruleum sp. nov. is pale grey
(Fig. 4C) while it is pink (Fig. 4D) in G. prasinum. In ad-
dition, the precloacal plate of Gonyosoma coeruleum sp.
nov. is divided (Fig. 4E) while it is single (Fig. 4F; Das
2013) in G. prasinum.
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137
Figure 6. Map showing the type locality of Gonyosoma coeruleum sp. nov. (blue star) in Mengla County, Yunnan Province, China;
the other collection sites of Gonyosoma coeruleum sp. nov. in Zhenyuan County (blue pentagon) and Menglian County (blue trian-
gle), Yunnan Province, China; the type locality (green square) of G. prasinum in Assam, India; and the new collection site (green
dot) of G. prasinum in Htamanthi wildlife sanctuary, Sagaing, Myanmar.
Figure 7. A habitat at the type locality of Gonyosoma coeruleum sp. nov. B Gonyosoma coeruleum sp. nov. asleep on a tree at night.
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Shuo Liu et al.: A new snake species from Chian138
Discussion
One type specimen of Gonyosoma prasinum is currently
available in the Natural History Museum, London (BMNH
1946.1.10.22). We have tried to check the type specimen but
failed due to the pandemic circumstances. However, we col-
lected two specimens from northwestern Myanmar, where
relatively close to the type locality, and these two specimens
agree with the descriptions and photographs (Das 2013;
Khandekar et al. 2021) of G. prasinum from the type locali-
ty and its vicinities, therefore we can conrm that the speci-
mens from northwestern Myanmar belongs to G. prasinum.
Although there is no major dierence in scalation be-
tween the new species and Gonyosoma prasinum in mor-
phological characters, however, the colorations of the iris
of them in life are obviously dierent. As, furthermore,
the new species and G. prasinum have a large genetic di-
vergence, they therefore deserve distinct a distinct status
at species level.
Previously, Gonyosoma prasinum has been recorded
in India, Bhutan, Myanmar, Thailand, Malaysia, Laos,
Vietnam, and China (Wangyal et al. 2020; Uetz et al.
2021), and within China, it was recorded from Yun-
nan, Guizhou, Sichuan, and Hainan Provinces (Zhao
et al. 1998; Wu et al. 2002; Zhao 2006; Yang and Rao
2008; Shi et al. 2011). However, we speculate that G.
prasinum is only distributed in India, Bhutan, northern
Myanmar, and Southeastern Tibet, China. Based on the
photographs (eg. Vogel and Pauwels 2004; Pauwels
et al. 2006; Zhao 2006; Ziegler et al. 2007; Yang and
Rao 2008; Shi et al. 2011; Hecht et al. 2013; Pham et
al. 2014; Qi 2019; Luu et al. 2020; Nguyen et al. 2020;
and unpublished data) of snakes previously identied as
G. prasinum from other parts of Yunnan Province, and
Sichuan, Guizhou, and Hainan Province, China, Thai-
land, Laos, and Vietnam, through the coloration of the
iris, we speculate that these populations also belong to
Gonyosoma coeruleum sp. nov. For the populations dis-
tributed in Malaysia, as we do not have specimens and
photographs, whether they also belong to Gonyosoma
coeruleum sp. nov. is still unknown.
Gonyosoma coeruleum sp. nov. is a familiar species in
southern Yunnan, China, however, we observed that Gon-
yosoma coeruleum sp. nov. was often captured and traded
as pets due to its beautiful appearance. In order to protect
this beautiful species, we suggest that this species to be
added to the local protected animal lists and be banned
from pet trading.
Acknowledgements
The eld survey in Myanmar was undertaken at the invi-
tation of the Republic of the Union of Myanmar, Ministry
of Natural Resources and Environmental Conservation,
Forest Department, Forest Research Institute. We thank
them for the invitation. Thanks also to the sta and the
local guides of Myanmar for their help in the eld. This
work was supported by Science-Technology Basic Con-
dition Platform from the Ministry of Science and Tech-
nology of the People’s Republic of China (Grant No.
2005DKA21402), the project of the second comprehen-
sive scientic investigation of Xishuangbanna National
Nature Reserve, and the project of Ministry of Ecology
and Environment of China: Investigation and assess-
ment of amphibians and reptiles in Jinghong City, Meng-
hai County, and Mengla County.
References
Bleeker P (1859) Gonyosoma jansenii Blkr., eene nieuwe slang van Ma-
nado. Natuurkundig Tijdschrift voor Nederlandsch Indië 16(4): e242.
Blyth E (1854) Notices and descriptions of various reptiles, new or lit-
tle known, Part 2. Journal of the Asiatic Society of Bengal 23(3):
287–302.
Boulenger GA (1894) Catalogue of the snakes in the British Museum
(Natural History). Volume II., containing the conclusion of the Col-
ubridæ Aglyphæ. British Museum Natural History, London, 382 pp.
Burbrink FT, Lawson R (2007) How and when did Old World rat snakes
disperse into the New World? Molecular Phylogenetics and Evolu-
tion 43(1): 173–189. https://doi.org/10.1016/j.ympev.2006.09.009
Burbrink FT, Lawson R, Slowinski JB (2000) Mitochondrial DNA
phylogeography of the polytypic North American rat snake
(Elaphe obsoleta): a critique of the subspecies concept. Evolution
54: 2107–2118. https://doi.org/10.1111/j.0014-3820.2000.tb01253.x
Chen X, Lemmon AR, Lemmon EM, Pyron AR, Burbrink FT (2017)
Using phylogenomics to understand the link between biogeographic
origins and regional diversication in ratsnakes. Molecular Phy-
logenetics and Evolution 111: 206–218. https://doi.org/10.1016/j.
ympev.2017.03.017
Chen X, McKelvy AD, Grismer LL, Matsui M, Nishikawa K, Bur-
brink FT (2014) The phylogenetic position and taxonomic status
of the Rainbow Tree Snake Gonyophis margaritatus (Peters, 1871)
(Squamata: Colubridae). Zootaxa 3881(6): 532–48. http://dx.doi.
org/10.11646/zootaxa.3881.6.3
Cox MJ, van Dijk PP, Nabhitabhata J, Thirakhupt K (1998) A Photo-
graphic Guide to Snakes and Other Reptiles of Thailand and South-
east Asia. Asia Books, Bangkok, 144 pp.
Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest 2: more
models, new heuristics and parallel computing. Nature Methods 9:
772–772. https://doi.org/10.1038/nmeth.2109
Das I (2002) A Photographic Guide to Snakes and other Reptiles of
India. New Holland Publishers, London, 144 pp.
Das I (2010) A Field Guide to the Reptiles of South-East Asia. New
Holland Publishers, London, 376 pp.
Das I (2012) A Naturalist’s Guide to the Snakes of South-East Asia –
Malaysia, Singapore, Thailand, Myanmar, Borneo, Sumatra, Java
and Bali. John Beaufoy Publishing, Oxford, 160 pp.
Das A (2013) Notes on ratsnakes from north-eastern India. In: Schulz
K-D (Eds) Old World ratsnakes – A collection of papers. Bushmaster
Publications, Würselen, 309–336.
Dowling HG (1951) A proposed standard system of counting ventral in
snakes. Journal of Herpetology 1(5): 97–99.
Gray JE (1853) Descriptions of some undescribed species of reptiles
collected by Dr. Joseph Hooker in the Khassia Mountains, East Ben-
Evolutionary Systematics 5 2021, 129–139
evolsyst.pensoft.net
139
gal, and Sikkim Himalaya. Annals and Magazine of Natural History
(2), 12(72): 386–392. https://doi.org/10.1080/03745485709495063
Hecht VL, Pham CT, Nguyen TT, Nguyen TQ, Bonkowski M, Ziegler
T (2013) First report on the herpetofauna of Tay Yen Tu Nature Re-
serve, northeastern Vietnam. Biodiversity Journal 4(4): 507–552.
Jablonski D, Kukushkin OV, Avcı A, Bunyatova S, Kumlutaş Y, Ilgaz Ç,
Polyakova E, Shiryaev K, Tuniyev B, Jandzik D (2019) The biogeog-
raphy of Elaphe sauromates (Pallas, 1814), with a description of a new
rat snake species. PeerJ 7: e6944. https://doi.org/10.7717/peerj.6944
Janssen HY, Pham CT, Ngo HT, Le MD, Nguyen TQ, Ziegler T (2019)
A new species of Lycodon Boie, 1826 (Serpentes, Colubridae) from
northern Vietnam. ZooKeys 875: 1–29. https://doi.org/10.3897/zoo-
keys.875.35933
Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS
(2017) ModelFinder: fast model selection for accurate phylogenet-
ic estimates. Nature Methods 14: 587–589. https://doi.org/10.1038/
nmeth.4285
Khandekar A, Roy P, Kunte K (2021) Reptiles of India, v. 1.25. https://
www.indianreptiles.org [Accessed on 25 March 2021]
Kumar S, Stecher G, Li M, Knyaz C, Tamura K (2018) MEGA X:
Molecular Evolutionary Genetics Analysis across computing plat-
forms. Molecular Biology and Evolution 35: 1547–1549. https://doi.
org/10.1093/molbev/msy096
Luu VQ, Dinh TS, Lo OV, Nguyen TQ, Ziegler T (2020) New records
and an updated list of reptiles from Ba Vi National Park, Vietnam.
Bonn zoological Bulletin 69(1): 1–9.
Mocquard F (1897) Notes herpétologiques. Bulletin du Muséum Na-
tional d’Histoire naturelle 3: 211–217. https://doi.org/10.5962/bhl.
part.19256
Nguyen TV, Brakels P, Maury N, Sudavanh S, Pawangkhanant P, Idiia-
tullina S, Lorphengsy S, Inkhavilay K, Suwannapoom C, Poyarkov
NA (2020) New herpetofaunal observations from Laos based on
photo records. Amphibian & Reptile Conservation 14(2): 218–249.
Pauwels OSG, Droz B, Vogel G (2006) Elaphe prasina (Blyth, 1854) in
Thailand. Elaphe 14(4): e62.
Peters W (1871) Über neue Reptilien aus Ostafrika und Sarawak (Bor-
neo), vorzüglich aus der Sammlung des Hrn. Marquis J. Doria zu
Genua. Monatsberichte der Königlichen Preussische Akademie Des
Wissenschaften zu Berlin 1871: 566–581.
Pham AV, Nguyen SLH, Nguyen TQ (2014) New records of snakes
(Squamata: Serpentes) from Son La Province, Vietnam. Herpetol-
ogy Notes 7: 771–777.
Purkayastha J (2013) An Amateur’s Guide to Reptiles of Assam. EBH
Publishers, Guwahati, 146 pp.
Qi S (2019) A photographic guide to reptiles of China. Chongqing Uni-
versity Press, Chongqing, 236 pp.
Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S,
Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2:
ecient Bayesian phylogenetic inference and model choice across
a large model space. Systematic Biology 61: 539–542. https://doi.
org/10.1093/sysbio/sys029
Schulz K-D (1996) A monograph of the colubrid snakes of the genus
Elaphe Fitzinger. Koeltz Scientic Books, Havlickuv Brod, 439 pp.
Schulz K-D (1991) Die hinterasiatischen kletternattern der Gattung
Elaphe Teil XX Elaphe prasina (Blyth 1854) und Elaphe frenata
(Gray 1853). Sauria 13(2): 15–17.
Shi HT, Zhao EM, Wang LJ, Bi H, Lyu SQ, Liu HN, Wang JC, Zhao H,
Hong ML (2011) Amphibian and Reptile Fauna of Hainan. Science
Press, Beijing, 285 pp.
Silvestro D, Michalak I (2012) raxmlGUI: a graphical front-end for
RAxML. Organisms Diversity and Evolution 12(4): 335–337.
https://doi.org/10.1007/s13127-011-0056-0
Smith MA (1943) The fauna of British India, Ceylon and Burma, in-
cluding the whole of the Indo-chinese subregion. Reptilia and Am-
phibia (Vol. III). Serpentes. Taylor and Francis, London, 583 pp.
Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins DG
(1997) The ClustalX windows interface: Flexible strategies for
multiple sequence alignment aided by quality analysis tools. Nu-
cleic Acids Research 25: 4876–4882. https://doi.org/10.1093/
nar/25.24.4876
Uetz P, Freed P, Hošek J (2021) The Reptile Database. http://www.rep-
tile-database.org [Accessed on 25 March 2021]
Vogel G, Pauwels OSG (2004) Geographic distribution. Elaphe prasina
(Green Bush Rat Snake). Herpetological Review 35(4): e410.
Wang K, Ren JL, Chen HM, Lyu ZT, Guo XG, Jiang K, Chen JM, Li
JT, Guo P, Wang YY, Che J (2020) The updated checklists of am-
phibians and reptiles of China. Biodiversity Science 28(2): 189–218.
http://www.biodiversity-science.net/CN/10.17520/biods.2019238
Wangyal JT, Bower DS, Sherub, Tshewang S, Wangdi D, Rinchen K,
Phuntsho S, Tashi C, Koirala BK, Gyeltshen, Bhandari GS, Jamtsho
S, Phuntsho Y, Koirala TP, Ghalley BB, Chaida L, Tenzin J, Powrel
RB, Tshewang R, Raika ON, Jamtsho S, Kinley, Gyeltshen, Tashi S,
Nidup D, Wanfdi N, Phuentsho, Norbu L, Wangdi K, Wangchuk T,
Tobgay P, Dorji T, Das I (2020) New Herpetofaunal Records from
the Kingdom of Bhutan Obtained through Citizen Science. Herpeto-
logical Review 51(4): 790–798.
Wu GJ, Wu TH, Zhu J, Xiao L (2002) A new record of snake species
in Sichuan Province – Elaphe prasina. Sichuan Journal of Zoology
21(2): 66.
Yang DT, Rao DQ (2008) Amphibia and Reptilia of Yunnan. Yunnan
Publishing Group Corporation, Yunnan Science and Technology
Press, Kunming, 411 pp.
Zhao EM (2006) Snakes of China. Anhui Science Technology Publish-
ing House, Hefei, 669 pp.
Zhao EM, Huang MH, Zong Y, Jiang YM, Huang QY, Zhao H, Ma JF,
Zheng J, Huang ZJ, Wei G, Yang DQ, Li DJ (1998) Fauna Sinica
Reptilia, vol. 3: Squamata: Serpentes. Science Press, Beijing, 522
pp.
Ziegler T, Hendrix R, Vu TN, Vogt M, Forster B, Dang KN (2007) The
diversity of a snake community in a karst forest ecosystem in the
central Truong Son, Vietnam, with an identication key. Zootaxa
1493: 1–40. https://doi.org/10.11646/zootaxa.1493.1.1
... This is the first record of this species in Vietnam. Gonyosoma coeruleum was recently described by Liu et al. (2021) from Yunnan Province, China. The new species closely resembles G. prasinum (Blyth), but it is differentiated from the latter species by having the precloacal plate divided, iris blue and inside of mouth greyish-white in life. ...
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