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343
A new species of the genus Amolops (Amphibia: Ranidae)
and the first national record of Amolops vitreus from
China
Yun-He Wu1,2,*, Zhong-Bin Yu1,2,*, Chen-Qi Lu1,3, Yin-Peng Zhang4, Wen-Jie Dong1,3, Xiao-Long
Liu5, Felista Kasyoka Kilunda1,3, Yun Xiong6, Yun-Fang Jiang7, Hong Ouyang7, Zhong-Xiong Fu8,
Yun-Biao He9, Zhi-Yong Yuan5, Jing Che1,2
1 State Key Laboratory of Genetic Resources and Evolution & Yunnan key laboratory of biodiversity and ecological conservation of Gaoligong
Mountain, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan 650223, China
2 Southeast Asia Biodiversity Research Institute, Chinese Academy of Sciences, Yezin, Nay Pyi Taw 05282, Myanmar
3 Kunming College of Life Science, University of the Chinese Academy of Sciences, Kunming, Yunnan 650204, China
4 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson 85721, USA
5 Key Laboratory of Freshwater Fish Reproduction and Development, Ministry of Education, Southwest University, Chongqing 400715, China
6 Gongshan Bureau of Gaoligongshan National Nature Reserve, Nujiang, Yunnan 673500, China
7 Lushui Bureau of Gaoligongshan National Nature Reserve, Nujiang, Yunnan 673299, China
8 Yunnan Senye Biotechnology Co., Ltd, Xishuangbanna, Yunnan 666100, China
9 Fugong Bureau of Gaoligongshan National Nature Reserve, Nujiang, Yunnan 673400, China
* Authors contributed equally to this work
https://zoobank.org/C01CD1E8-38F6-4908-93CD-0F7C8259618B
Corresponding authors: Jing Che (chej@mail.kiz.ac.cn), Zhi-Yong Yuan (yuanzhiyongkiz@126.com)
Academic editors Raael Ernst / Martin Päckert | Received 14 June 2023 | Accepted 09 January 2024 | Published 04 April 2024
Citation: Wu Y-H, Yu Z-B, Lu C-Q, Zhang Y-P, Dong W-J, Liu X-L, Kilunda FK, Xiong Y, Jiang Y-F, Ouyang H, Fu Z-X, He Y-B, Yuan Z-Y,
Che J (2024) A new species of the genus Amolops (Amphibia: Ranidae) and the rst national record of Amolops vitreus from China. Vertebrate
Zoology 74 343–357. https://doi.org/10.3897/vz.74.e108013
Abstract
The torrent frogs of the genus Amolops represent a great anuran diversication in southern China and Southeast Asia. Previous
studies have shown that, the diversity of this genus still remains underestimated. During herpetological surveys from 2021 to 2022,
several Amolops specimens were collected from the international border regions of southwestern Yunnan Province, China. Herein,
we utilized molecular phylogenetic and morphological data to identify these specimens. Our ndings indicate the presence of a sepa-
rate and previously unknown lineage in the A. viridimaculatus group, which we formally describe as a new species. Furthermore, the
specimen from Xishuangbanna National Nature Reserve clustered with A. vitreus from the paratype, supporting the morphological
diagnosis. Therefore, we describe a new species and a new species record for China. Our study contributes to the species richness
of the genus Amolops as well as the diversity of amphibians in China. Notably, our discovery brings the total number of Amolops
species to 85 and the total number of torrent frog species known to occur in China to 53. In addition, our study further conrmed
that Yunnan and Indochina Peninsula have similar faunal composition, implying that more studies are needed to achieve a complete
understanding of the species diversity and distribution pattern.
Key words
Amolops yangi sp. nov., Amolops vitreus, new species, new species record, Southwest border of China, Yunnan Province
Vertebrate Zoology 74, 2024, 343–357 | DOI 10.3897/vz.74.e108013
Copyright Yun-He Wu et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution,
and reproduction in any medium, provided the original author and source are credited.
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
344
Introduction
Yunnan is located at the intersection of three global bio-
diversity hotspots: the eastern Himalayas region, the
mountains of Southwest China, and the Indo-Burma
region (Myers et al. 2000). It is recognized globally as
one of the richest and most diverse regions due to its spe-
cies richness and wide range of habitats and ecosystems
(Yang et al. 2004; Pu et al. 2007). Although Yunnan only
comprises 4.1% of China’s total land area, it is home to
more than 19,365 higher plant and 2,273 vertebrate spe-
cies (54.8% of China’s total) (Liu et al. 2021). Moreover,
this area harbors more than 3,400 endemic species and
about 60% of the endangered species in China (Yang et
al. 2004; Liu et al. 2021). Notably, current research still
indicates that the vertebrate diversity of Yunnan has been
underestimated (e.g., Chen et al. 2015; Su et al. 2020;
Wang et al. 2022). The gap may be even larger in inver-
tebrates (e.g., Yu et al. 2022). In particular, it shares an
extensive 4,060-kilometer border with Vietnam, Laos and
Myanmar. Since, these areas are located in the same zoo-
geographic region (Holt et al. 2013), and their habitats on
either side of the border are continuous with no known
biogeographic barriers, their species composition is sim-
ilar. Furthermore, many new records have been reported
in Yunnan in recent years (e.g., Jiang et al. 2022; Yu et al.
2022). Similarly, many new records have been reported
from Vietnam, Laos and Myanmar, for species initially
described from China (e.g., Nguyen et al. 2020; Rahman
et al. 2020; Liang et al. 2023).
Amphibians face a greater risk of rapid loss of di-
versity and are considered the most threatened group of
vertebrates (Alroy 2015). Yunnan Province is the region
with the richest amphibian diversity in China, and the
number of threatened amphibian species has increased
signicantly in recent years (Yuan et al. 2022). There-
fore, a clear number and distribution of species is both
warranted and of great conservation importance. Recent
studies on Megophrys sensu lato, Leptobrachella, and
Amolops have shown that the diversity of amphibians in
this region is underestimated (Chen et al. 2017, 2018; Wu
et al. 2020). Intensied eldwork surveys in recent years
have led to a continuous description of new amphibian
species (Chen et al. 2020; Liu et al. 2021, 2023; Shi et al.
2021; Wu et al. 2019; Yang et al. 2019a,b). Furthermore,
a series of new genera records and new species records
of amphibians from China have been described along the
border region in the recent past, such as the genus Wija-
yarana (Wu et al. 2023a), Nidirana chapaensis (Yuan et
al. 2018), Amolops putaoensis (Zhang et al. 2022), and
Microhyla hmongorum (Wu et al. 2023b). These ndings
highlight a largely underestimated amphibian diversity in
the border region of Yunnan Province.
The genus Amolops Cope, 1865 of the family Rani-
dae is the most species-rich genus within the family Ra-
nidae, distributed widely throughout Nepal, northern In-
dia, western and southern China to Malay Peninsula, and
currently includes 84 recognized species (Frost 2023).
Amolops is characterized by possessing a gastromyzo-
phorus adhesive disk (ventral sucker), dorsal and ventral
poison glands in their tadpoles and enlarged digital discs
in adults (Yang 1991; Fei et al. 2009; Wu et al. 2020).
These montane species mainly inhabit fast-flowing, rocky
stream environments, such as torrents and waterfalls (Fei
et al. 2009; Wu et al. 2020). In recent years, extensive
taxonomic and phylogenetic studies have been conducted
on the genus Amolops, delimiting 10 species groups (e.g.,
Wu et al. 2020; Zeng et al. 2020; Jiang et al. 2021), which
greatly improves our understanding on the taxonomy and
species diversity of this genus. In China, 51 species have
been recorded, and assigned into eight species groups:
A. chayuensis group, A. daiyunensis group, A. hainan-
ensis group, A. mantzorum group, A. marmoratus group,
A. monticola group, A. ricketti group and A. viridimac-
ulatus group (AmphibiaChina 2023). Nearly half of the
currently recognized Chinese Amolops species have been
described in the last 10 years (AmphibiaChina 2023).
Moreover, 22 species of Amolops have been recorded in
Yunnan, 13 of which were rst discovered in Yunnan in
the past 10 years (AmphibiaChina 2023). This is ascribed
to intensied survey eorts and the adoption of more in-
tegrative taxonomic approaches.
During our recent herpetological surveys at the Yun-
nan border area, four specimens of the genus Amolops
were collected. Molecular data and morphological com-
parisons revealed that these specimens included a new
species and a new species record for China which we
herein describe.
Materials and Methods
Sampling
A total of four individuals including three adult males and
one adult female were collected from the Yunnan border,
China, (Fig. 1). Following euthanasia, all specimens were
xed in 10% formalin for 24 hours for preservation after
extraction of liver tissues from the adults in 95% ethanol,
and subsequently transferred to 70% ethanol for perma-
nent preservation. Voucher specimens were deposited in
the herpetological collection of the Museum of the Kun-
ming Institute of Zoology (KIZ), Chinese Academy of
Sciences (CAS).
DNA extraction, PCR amplification,
and sequencing
Total genomic DNA was extracted using standard phe-
nol-chloroform protocols (Sambrook et al. 1989), and three
partial fragments of the mitochondrial 16S rRNA gene
(16S), cytochrome oxidase subunit I (COI), and NADH
dehydrogenase subunit 2 (ND2) genes were amplied and
sequenced using the following primers: 16SAR (5’-CGC-
CTGTTTAYCAAAAACAT-3’) and 16SBR (5’-CC G-
G T YTG AACTCAGATCAYGT-3) (Kocher et al. 1989),
Vertebrate Zoology 74, 2024, 343–357 345
ND2AR (5’-CAATGTTGGTTAAA AT C CTTCC-3’) and
NDBR (5’-AGGCTTTGAAGGCC TTTGGTC-3’) (Stu-
art et al. 2006), and Chmf4 (5’-TYTCWACWAAYCA Y-
A A A GAYATCGG-3’), and Chmr4 (5’-ACYTCRGGR-
T G R C CRAARAATCA-3’) (Che et al. 2012). The
poly merase chain reaction (PCR) was performed in a 25
µl reaction volume with the following cycling conditions:
initial denaturing step at 95°C for 4 min, 35 cycles of de-
naturing at 94°C for 40 s, annealing at 55°C for 16S rRNA
for 45 s, 50°C for ND2 for 1 min, and 55°C for COI for
45 s, extending at 72°C for 1 min and a nal extension at
72°C for 10 min. PCR products were directly sequenced
by an ABI 3730xl DNA automated sequencer (Applied
Biosystems, UK) with both forward and reverse primers.
All sequences were assembled from forward and reverse
reads and edited manually using DNASTAR LASER-
GENE 7.1. All new sequences were deposited in Gen-
Bank (Table S1).
Phylogenetic analysis
To study the historical relationships among Amolops spe-
cies, phylogenetic trees were reconstructed based on the
16S, COI, and ND2 fragments. Homologous sequences
of Amolops and representative outgroups (A. spinapec-
toralis and A. yatseni) were downloaded from GenBank
(Table S1). Phylogenetic relationships were inferred us-
ing maximum likelihood (ML) and Bayesian inference
(BI) methods. The best-t nucleotide substitution model
was selected using the JMODELTEST v2.1.7 (Darriba
et al. 2012) following the Bayesian information criteri-
on (BIC; Posada 2008) for BI. The best-t models were
summarized in Table S2. BI analysis was implemented
by the CIPRES web server (Miller et al. 2010). For BI
analyses, the Monte Carlo Markov chain length was run,
sampling one tree every 1,000 generations for 10,000,000
generations with a burn-in of 25%. Convergence was
assessed in TRACER 1.5 (Rambaut et al. 2009) by the
average standard deviation of split frequencies (below
0.01) and the eective sample size (over 200). Maximum
likelihood (ML) analyses were conducted using RAxML
v8.0.0 with 1,000 bootstrap replicates and using the stan-
dard bootstrap search (random seed value 12,345) under
GTR + I + G nucleotide substitution model (Stamatakis et
al. 2014). We also calculated pairwise sequence diver-
gence using uncorrected pairwise distances (p distances)
implemented in MEGA 6.0.6 (Tamura et al. 2013).
Morphology
The four preserved adult specimens were measured with
digital calipers to the nearest 0.1 mm. Measurements fol-
lowed Fei et al. (2009). Measurements included the fol-
lowing: SVL (Snout-vent length): measured from tip of
snout to vent; HL (head length): measured from tip of
snout to jaw angle; HW (head width): measured as head
width at its widest point; SL (snout length): measured
from tip of snout to anterior corner of eye; INS (inter-
nasal space): measured as distance between nares; IOS
(interorbital space): measured at narrowest point between
eyelids on top of head; NED (nasal to eye distance): mea-
sured as distance from the anterior corner of eye to nostril
center; UEW (upper eyelid width): maximum width of
upper eyelid; ED (eye diameter): measured as the distance
between corners of eye; TD (tympanum diameter): mea-
sured as maximal diameter of tympanum; LAHL (length
of lower arm and hand): distance from elbow to the tip
of the third nger; HND (hand length): measured as the
distance from the proximal edge of inner metacarpal tu-
bercle to the tip of third nger; LAD: (diameter of lower
arm); FEM (femoral length): measured from the cloaca
to the knee; TIB (tibia length): measured as the distance
from knee to heel; FTL (foot length): measured as the
distance from proximal end of inner metatarsal tubercle
to the tip of fourth toe. Sexual maturity was determined
Figure 1. Sampling localities of
Amolops used in this study. The
green pentagram indicates the type
locality of Amolops yangi sp. nov.,
the green circles indicate the other
distribution localities of Amolops
yangi sp. nov., the red pentagram
indicates the type locality of A.
vitreus, and the red circle indicates
the new locality record of A. vit-
reus.
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
346
by the presence of nuptial pads or vocal sacs (males), or
eggs or enlarged oviducts (females).
To compare the morphometrics of A. viridimaculatus
group, we extracted data from the published literature for
all species available (Table S3). Principal Component
Analysis (PCA) was used to explore the morphological
dierences between the undescribed species and other
species of A. viridimaculatus group. The morphometric
analyses were conducted separately for the male and fe-
male groups. The analyses were carried out in R 4.0.2.
Figure 2. Phylogram of Amolops based on mitochondrial 16S, COI, and ND2. The “*” denote Bayesian posterior probabilities
(BPP) = 1.00 and bootstrap support (BS) = 100%; “#” denote Bayesian posterior probabilities (BPP) ≥ 0.95 and bootstrap support
(BS) ≥ 70%. Node values with Bayesian posterior probabilities (BPP) < 95 or Bootstrap support (BS) < 70 are not shown.
Vertebrate Zoology 74, 2024, 343–357 347
Results
Phylogenetic data
Our concatenated mtDNA alignment (ND2: 890 base
pairs (bp), COI: 570 bp, 16S: 541 bp) contained 108 indi-
viduals with a total of 2002 bp, with 900 conserved sites
and 1098 variable sites. Of the variable sites in the align-
ment, 1008 were parsimony-informative (559 bp, 247 bp,
and 202 bp including the outgroup sequences).
Both BI and ML analyses resulted in essentially iden-
tical topologies, with relatively robust support for most
nodes. The genus Amolops was recovered as monophylet-
ic with strong support from both analyses (BPP = 1; BS =
96; Fig. 2). The genus forms nine monophyletic clades
(clade I–IX, Fig. 2), corresponding to nine dierent spe-
cies groups. Among these, three collected specimens from
Fugong and Lushui formed a separate lineage with strong
support (BPP = 1; BS = 100; Fig. 2) and nested within
the A. viridimaculatus group (clade III). The genetic dis-
tance between the new population and other species of
the A. viridimaculatus group ranged from 2.3% (with A.
kaulbacki) to 3.5% (with A. beibengensis) for 16S, 8.3%
(with A. medogensis) to 11.1% (with A. beibengensis)
for COI, and 9.4% (with A. formosus) to 12.9% (with A.
kaulbacki) for ND2 (Table S4). It is comparable to the di-
vergences among the nearest neighbour genetic distances
of this group, which ranged from 1.0% (A. wangyali and
A. tawang) to 6.1% (A. himalayanus and A. wangyufani)
for 16S, 6.1% (A. viridimaculatus and A. kaulbacki) to
10.2% (A. beibengensis and A. formosus) for COI, and
6.6% (A. viridimaculatus and A. beibengensis) to 14.3%
(A. kaulbacki and A. formosus) for ND2 (Table S4). Mor-
phologically, the new specimens diered from present-
ly recognized congeners. In the PCA analysis, the rst
two principal components explained 85.7% of the total
variation in the males, where PC1 and PC2 eigenvectors
accounted for 75.1% and 10.7% of the total variance, re-
spectively (Table S5). Similarly, the rst two principal
components occupied a considerable proportion in the
females, 80.9% of the total, whereas PC1 and PC2 eigen-
vectors accounted for 65.8% and 15.1% of the total vari-
ance, respectively (Table S5). Regardless of the sex of the
species, the samples showed no overlap between the new
specimens and other species of A. viridimaculatus group
on the two-dimensional graphs for PC1 and PC2 (Fig. 3).
In addition, one collected male specimen (KIZ 050452)
from Xishuangbanna National Nature Reserve was nest-
ed in the A. monticola group (Fig. 2, clade I) and formed
a monophyletic clade with A. vitreus from the paratype
conrming shared genetic traits therefore validating the
morphological diagnosis. As a result, below we describe
a new species and a new species record for China.
Taxonomic account
Amolops yangi Wu, Yu, Lu, Yuan & Che
sp. nov.
https://zoobank.org/5B4E8AC7-37AD-4192-83E3-
A05E49393B7B
Holotype. Adult female (KIZ 038643), from Ega, Lushui,
Nujiang Lisu Autonomous Prefecture, Yunnan Prov-
ince, China (26.43744°N, 98.75044°E; elevation 3496
m a.s.l.), collected by Zhong-Bin Yu, Dong An, Tian-En
Chen on 07, August, 2021.
Paratypes. One adult male KIZ 038645 from Ega,
Lushui, Nujiang Lisu Autonomous Prefecture, Yunnan
Province, China (26.44924°N, 98.76762°E; elevation
2915 m a.s.l.), collected by Zhong-Bin Yu, Dong An,
Tian-En Chen on 07, August, 2021; one adult male KIZ
050788 from Yaping road, Lumadeng Township, Fugong
county, Nujiang Lisu Autonomous Prefecture, Yunnan
Province, China (27.15890°N, 98.79712°E; elevation
2451 m a.s.l.), collected by Zhong-Bin Yu, Dong An on
05, August, 2022.
Etymology. The specic epithet “yangi” is a patronymic
noun in the genitive singular; derived from the name of
Prof. Da-Tong Yang of the Kunming Institute of Zoolo-
gy, CAS, China. We acknowledge his great contributions
Figure 3. Plots of the rst principal component (PC1) versus the second (PC2) for A. yangi sp. nov. and other species of A. viridi-
maculatus group from the principal component analysis. A Male, B female.
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
348
to the herpetological research in southwestern China. We
suggest the Chinese formal name as “杨氏湍蛙”.
Diagnosis. Amolops yangi sp. nov. is assigned to the ge-
nus Amolops based on molecular phylogenetic analyses
and can be distinguished from its congeners by a com-
bination of the following characters: (1) medium body
size (SVL 46.3–51.8 mm in males and at least 51.5 mm
in female); (2) vomerine teeth developed, on two short
oblique between choanae, equal in distance from each
other as to choanae; (3) supernumerary tubercles pres-
ent at the base of each nger; (4) tympanum indistinct;
(5) three metacarpal tubercles, inner metacarpal tubercle
long, outer metacarpal tubercle relatively small, oval,
median one rounded; (6) supratympanic fold indistinct;
(7) discontinuous glandular dorsolateral fold from rear of
eye to near vent; (8) circummarginal grooves present on
tips of outer three ngers, absent on rst nger; (9) iris
distinctly bicolored, golden-yellow in upper one-fourth
and reddish brown in lower three-fourths, black reticu-
lations throughout; (10) rictal gland absent; (11) dorsal
surface of the head, back, limbs, ngers, and toes green,
interspersed with irregular black spots; (12) dorsal parts
of limbs, ngers and toes with black crossbars; (13) vocal
sac absent in males; (14) male with orange nuptial pad at
the base of rst nger.
Description of holotype (all measurements in mm; see
Table 1). KIZ 038643, sexually mature female, body size
moderate, adult female (SVL 51.5 mm); head length larger
than wide (HL/SVL 34.2%, HW/SVL 32.6%); top of head
flat; snout short (SL/HL 46.0%), snout rounded in dorsal
view (Fig. 4A), obtusely rounded in prole, projecting
beyond margin of lower jaw; loreal region and concave
and oblique; canthus rostralis distinct, slightly constricted
behind nostrils; dorsal region of snout flattened; eyes rel-
atively large (ED/HL 29.0%), slightly protuberant in dor-
sal view and notably protruding in prole (Fig. 4A), eye
diameter shorter than snout length (ED/SL 63.0%); nos-
trils oval, laterally orientated, slightly protuberant, closer
to anterior corner of eye than to tip of snout; pupil oval,
horizontal; tympanum indistinct, circular in shape, rela-
Table 1. Measurements (mm) of Amolops yangi sp. nov. and A. vitreus. The asterisk (*) indicates the holotype.
A. yangi sp. nov. A. yangi sp. nov. A. yangi sp. nov.* A. vitreus
Sex ♂ ♂ ♀ ♂
Catalog No. KIZ 050788 KIZ 038645 KIZ 038643 KIZ 050452
SVL 51.8 46.3 51.5 38.9
HL 17.0 16.8 17.6 15.4
HL/SVL 32.8% 36.3% 34.2% 39.6%
HW 16.7 15.4 16.8 13.9
HW/SVL 32.2% 33.3% 32.6% 35.7%
SL 7.2 7.7 8.1 6.5
SL/SVL 13.9% 16.6% 15.7% 16.7%
SL/HL 42.4% 45.8% 46.0% 42.2%
INS 6.7 6.5 6.9 4.6
INS/SVL 12.9% 14.0% 13.4% 11.8%
IOS 4.1 4.8 4.3 4.1
INS/IOS 163.4% 135.4% 160.5% 112.2%
NED 3.3 3.2 3.3 3.4
UEW 4.1 3.6 4.1 3.8
INS/UEW 163.4% 180.6 168.3% 121.1%
UEW/IOS 100.0% 75.0% 95.3% 92.7%
ED 5.0 4.9 5.1 4.4
ED/HL 29.4% 29.2% 29.0% 28.6%
ED/SL 69.4% 63.6% 63.0% 67.7%
TD 2.6 2.4 1.8 2.7
TD/ED 52.0% 49.0% 35.3% 61.4%
LAHL 26.9 25.0 29.1 19.6
LAHL/SVL 51.9% 54.0% 56.5% 50.4%
HND 17.1 15.4 18.4 12.2
HND/SVL 33.0% 33.3% 35.7% 31.4%
LAD 5.0 4.8 4.4 3.8
LAD/SVL 9.7% 10.4% 8.5% 9.8%
FEM 26.3 22.7 25.3 22.2
TIB 27.1 24.3 27.2 23.5
FEM/TIB 97.0% 93.4% 93.0% 94.5%
FTL 27.7 26.8 28.0 20.3
FEM/FTL 94.9% 84.7% 90.4% 109.4%
Vertebrate Zoology 74, 2024, 343–357 349
tively small (TD/HL 10.2%), tympanum diameter about
one third of eye diameter (TD/ED 35.3%); internarial dis-
tance (INS/SVL 13.4%) larger than width of upper eyelid
(INS/UEW 168.3%) and interorbital distance (INS/IOS
160.5%); tongue cordiform, deeply notched posteriorly;
vomerine teeth developed, on two short oblique between
choanae, equal in distance from each other as to choanae;
choanae oval; maxillary teeth developed; a small tooth-
like projection on anteromedial edge of mandible.
Forelimbs moderately long and robust, forelimb and
hand length (29.1 mm) longer than half body size (LAHL/
SVL 56.5%); relative length of ngers: I<II<IV<III; cir-
cummarginal grooves present on tips of outer three n-
gers, absent on rst nger; subarticular tubercles promi-
nent and oval, formula 1, 1, 2, 2; supernumerary tubercles
present at the base of each nger; webbing between n-
gers absent; narrow lateral fringes of ngers III and IV;
three metacarpal tubercles, inner metacarpal tubercle
long, outer metacarpal tubercle relatively small, oval,
median one rounded (Fig. 4D).
Hindlimbs long and robust, femoral length shorter than
the tibia length (FEM/TIB 93.0%) and the foot length
(FEM/FTL 90.4%); tibiotarsal articulation of adpressed
limb reaching between nostrils and eyes when hindlimb
stretched alongside of body; the heels overlapping when
the tibias are perpendicular to the body axis; relative toes
lengths: I<II<III<V<IV; narrow lateral fringes of preaxial
side of toe I and postaxial side of toe V; tips of all toes
Figure 4. Amolops yangi sp. nov. (Holotype KIZ 038643). A Lateral view, B dorsal view, C ventral view of thighs, D ventral view
of hand, E foot, F habitat. Photos by Zhong-Bin Yu.
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
350
expanded into discs with circummarginal grooves; toes
fully webbed except for fourth toe, in which web reaches
beyond distal subarticular tubercle; subarticular tubercles
oval and distinct, formula 1, 1, 2, 3, 2; supernumerary
tubercles absent; inner metatarsal tubercle long, outer
metatarsal tubercle absent (Fig. 4E).
Dorsal surface of head, body, limbs, ngers, toes and
flank of body relatively smooth; loreal region densely
scattered with raised tubercles; temporal region and pos-
terior angle of the jaw with dense tubercles; skin ventral-
ly smooth, including throat, chest, abdomen, and ventral
surface of limbs; discontinuous glandular dorsolateral
fold from rear of eye to near vent; supratympanic fold
indistinct; rictal gland absent (Fig. 4A–E).
Color of holotype in life. For coloration of the holotype
in life see Figure 4A-E. Dorsal surface of the head, back,
limbs, ngers, and toes green, interspersed with irregular
black spots; throat, ventral surface of the head, chest and
anterior abdomen mostly yellow, with scattered grayish
spots; upper part of flanks green, lower part of flanks
green-yellow; a black stripe below edge of the canthus
rostralis extending from the snout tip across the eyes, to
the anterior edge of supratympanic fold; upper lips with
three dark bars; ventral surface of thighs orangish, dense-
ly scattered with small yellow spots; dorsal parts of limbs,
ngers and toes with black crossbars; ventral surface of
ngers and toes orange; toes webbing yellowish-gray;
the inside of lower arm with black stripe; ventral surface
of all ngers discs, subarticular tubercles, supernumer-
ary tubercles, and metacarpal tubercles orange, ventral
surface of outer three toes discs, subarticular tubercles,
and metatarsal tubercle grey; ventral surface of inner two
toes discs orange; iris distinctly bicolored, golden-yellow
in upper one-fourth and reddish brown in lower three-
fourths, black reticulations throughout.
Color of holotype in preservative. For coloration of the
holotype in preservative see Fig. 5A–D. After two years
of storage in ethanol, the dorsal surface fading to metallic
blue with irregular black spots; black crossbars present
on dorsal surfaces of limbs, ngers and toes still clear;
throat, chest, belly, and ventral surface of limbs fading to
cream-yellow, with irregular gray pigmentations; ventral
surface of the hands and toes cream-yellow, digit tips and
subarticular tubercles fading to cream-yellow or gray-
ish-white.
Male secondary sexual characteristics. Adult males
possess orange nuptial pads covering the base of rst n-
ger; absence of vocal sacs in males.
Distribution and ecology. Amolops yangi sp. nov. is
currently known from two localities in the Gaoligong
Mountains. These are Fugong and Lushui County, both
in Yunnan Province, China. These two localities are sepa-
rated by a straight-line distance of approximately 80 km.
The new species inhabits the banks of rocky, fast-flow-
ing streams or perches on shrubs (ca. 0.5 m above the
ground) along the swift flowing streams (Fig. 4F). The
Figure 5. A Dorsal view, B ventral view, C ventral view of hand, D ventral view of foot, (D) Amolops yangi sp. nov. ( Holotype KIZ
038643) in preservative. Photos by Zhong-Bin Yu.
Vertebrate Zoology 74, 2024, 343–357 351
new species primarily inhabits high altitude mountainous
areas with elevations ranging from 2500 to 3500 m. The
breeding season is currently uncertain. Other sympatric
amphibian species found in the same habitat included
Nanorana chayuensis, A. viridimaculatus, Scutiger gong-
shanensis, and Xenophrys glandulosa.
Comparisons. Phylogenetic analyses indicated that the
new species belongs to the A. viridimaculatus group
with strong support. Geographically, the new species is
found in Fugong and Lushui, Yunnan Province, China
that belong to Gaoligong Mountains, and close to north-
ern Myanmar. Therefore, we compared Amolops yangi
sp. nov. with morphologically, geographically, and mo-
lecularly similar species, which include A. chayuensis,
A. bellulus, A. putaoensis, A. binchachaensis, A. deng,
A. jin jiangensis, A. viridimaculatus, A. kaulbacki, A. mar-
moratus, A. afghanus, and A. tuberodepressus, A. bei-
ben gensis, A. wangyufani, A. formosus, A. medogensis,
A. pallasitatus, A. nidorbellus, A. himalayanus, A. wang-
yali, A. longimanus, A. ailao, A. chanakya, and A. tawang
(Andersson 1938; Jiang et al. 1983; Yang and Rao 2008;
Fei et al. 2009; Biju et al. 2010; Dever et al. 2012; Sun
et al. 2013; Nidup et al. 2016; Qi et al. 2019; Che et al.
2020; Gan et al. 2020; Liu and Yang 2000; Liu et al. 2000;
Mahony et al. 2022; Rao et al. 2022; Saikia et al. 2022;
Tang et al. 2023).
Amolops yangi sp. nov. is signicantly dierent from
A. viridimaculatus by discontinuous glandular dorsolat-
eral fold (vs. absent), supratympanic fold indistinct (vs.
distinct), SVL 46.3–51.8 mm in males and 51.5 mm in
female (vs. 72.7–82.3 in males and 83.0–94.3 in females),
dorsal surface of the head, back, limbs, ngers, and toes
green, interspersed with irregular black spots (vs. dorsum
and flank with nearly round green or yellowish green
spots, scattered with small green spots); from A. kaulbac-
ki by SVL 46.3–51.8 mm in males and 51.5 mm in female
(vs. SVL 72.6–82.6 mm in males and 82.7–87.2 mm in
females), discontinuous glandular dorsolateral fold (vs.
absent), supratympanic fold indistinct (vs. distinct), iris
distinctly bicolored, golden-yellow in upper one-fourth
and reddish brown in lower three-fourths, black reticula-
tions throughout (vs. eyes brownish black with scattered
yellow spotting and yellow ring around iris); from A.
beibengensis by SVL 46.3–51.8 mm in males and 51.5
mm in female (vs. SVL 75.8 mm males and 90.2–93.2
mm in females), supratympanic fold indistinct (vs. dis-
tinct, wide and thick), discontinuous glandular dorsolat-
eral fold from rear of eye to near vent (vs. absent); from A.
wangyufani by SVL 46.3–51.8 mm in males and 51.5 mm
in female (vs. SVL 68.3–69.0 mm males and 83.4 mm in
female), vomerine teeth developed, on two short oblique
between choanae, equal in distance from each other as
to choanae (vs. vomerine teeth developed, the two rows
are almost in touch), discontinuous glandular dorsolateral
fold from rear of eye to near vent (vs. absent); from A. me-
dogensis by SVL 46.3–51.8 mm in males and 51.5 mm in
female (vs. SVL 95.0 mm male and 72.4–96.9 mm in fe-
males), tibiotarsal articulation of adpressed limb reaching
between nostrils and eyes (vs. beyond tip of snout), su-
pratympanic fold indistinct (vs. distinct, wide and thick);
from A. himalayanus by dorsal surface of the head, back,
limbs, ngers, and toes green, interspersed with irregular
black spots (vs. dark brown, interspersed with irregular
yellow spots), vocal sac absent in males (vs. externally
visible vocal sacs present); from A. pallasitatus by SVL
51.5 mm in female (vs. SVL 70.6–72.3 mm in females),
discontinuous glandular dorsolateral fold from rear of eye
to near vent (vs. absent), dorsal surface of the head, back,
limbs, ngers, and toes green, interspersed with irregular
black spots (vs. dorsum yellow-green, with irregular dark
brown blotches without margins); from A. wangyali by
SVL 46.3–51.8 mm in males and 51.5 mm in female (vs.
SVL 71.4–76.7 mm males and 80.5–89.6 mm in females),
dorsal surface of the head, back, limbs, ngers, and toes
green, interspersed with irregular black spots (vs. large
brown irregularly shaped blotches on dorsum of head and
body), rictal gland absent (vs. a distinct patch of rictal
glands at rear of jaw on either side); from A. nidorbellus
by SVL 46.3–51.8 mm in males and 51.5 mm in female
(vs. SVL 76.4–82.3 mm males and 85.4–98.0 mm in fe-
males), dorsum green, interspersed with irregular black
spots (vs. dorsally brown with small irregularly arranged
cobalt green spots), discontinuous glandular dorsolateral
fold (vs. absent); from A. longimanus by SVL 46.3–51.8
mm in males and 51.5 mm in female (vs. SVL 30 mm),
nostrils closer to anterior corner of eye than to tip of snout
(vs. nostrils a litter nearer to tip of snout than the eye),
above tympanum to the forelimb a thick glandular paro-
toid-like swelling absent (vs. present), eye diameter (ED/
SL 63.0%) shorter than snout length (vs. snout about as
long as the eye diameter, ED/SL 97.7%); from A. formo-
sus by dorsal parts of limbs, ngers and toes with black
crossbars (vs. legs and toes with black white-dotted cross-
bars), inner metatarsal tubercle long, outer metatarsal tu-
bercle absent (vs. metatarsal tubercle indistinct), dorsum
green, interspersed with irregular black spots (vs. upper
parts green, marbled with black, the black spots enclosing
a number of small whitish dots); from A. chanakya by
SVL 46.3–51.8 mm in males (vs. SVL 76.4 mm), tym-
panum indistinct (vs. tympanum distinct, about 40% of
eye length), absence of vocal sacs in males (vs. vocal sac
externally visible), dorsal color green, interspersed with
irregular black spots (vs. dorsal color dull brick-red, spot-
ted with irregular cocoa-brown spots, these cocoa-brown
spots enclosing a number of smaller dull brick-red spots);
from A. tawang by SVL 46.3–51.8 mm in males (vs. SVL
82.5 mm), tibiotarsal articulation of adpressed limb reach-
ing between nostrils and eyes (vs. reaching up to snout),
dorsal color green, interspersed with irregular black spots
(vs. dorsal color olivegreen, spotted with large, irregular
shaped dark-brown spots, brown spots enclosing a num-
ber of small olivegreen dots).
Amolops yangi sp. nov. is signicantly dierent from
A. chayuensis by absence of vocal sacs in males (vs. pair
of external subgular vocal sacs); discontinuous glandular
dorsolateral fold from rear of eye to near vent (vs. dorso-
lateral fold prominent); upper part of flanks green, low-
er part of flanks green-yellow (vs. upper part of flanks
brown, lower part of flanks light green or white with dark
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
352
brown blotches); from A. bellulus by tympanum indis-
tinct (vs. tympanum distinct), iris distinctly bicolored,
golden-yellow in upper one-fourth and reddish brown
in lower three-fourths, black reticulations throughout
(vs. upper half of iris golden yellow with some irregular
brown spots, lower half dark brown); from A. putaoensis
by SVL of adult male 46.3–51.8 mm (vs. 37.6–40.2 mm),
rictal gland absent (vs. two rictal glands present), iris dis-
tinctly bicolored, golden-yellow in upper one-fourth and
reddish brown in lower three-fourths, black reticulations
throughout (vs. upper one-fourth of iris bronze with black
reticulations, lower three-fourths dark), absence of vocal
sacs in males (vs. pair of internal subgular vocal sacs
present); from A. binchachaensis by SVL of adult female
51.5 mm (vs. 65.0 mm), tympanum indistinct, relative-
ly small, about one third of eye diameter (vs. tympanum
big), supratympanic fold indistinct (vs. supratympanic
fold absent), dorsal surface of the head, back, limbs, n-
gers, and toes green (vs. light yellow); from A. deng by
SVL of adult female 51.5 mm (vs. 68.5–72.0 mm), tym-
panum indistinct, relatively small, about one third of eye
diameter (vs. tympanum distinct, slightly less than half of
eye diameter), tibiotarsal articulation of adpressed limb
reaching between nostrils and eyes (vs. tibiotarsal artic-
ulation of adpressed limb reaching beyond tip of snout);
from A. jinjiangensis by supratympanic fold indistinct
(vs. distinct), dorsal surface of head, body, limbs, n-
gers, toes and flank of body relatively smooth (vs. rough
with tubercles), the absence of a pair of large tubercles
on sides of cloaca (vs. present); from A. tuberodepres-
sus by supratympanic fold indistinct (vs. present, wide),
flanks smooth (vs. with flatter tubercles), relative length
of ngers I<II<IV<III (vs. II<IV<I<III), vomerine teeth
developed (vs. weak); from A. ailao by SVL 46.3–51.8
mm in males (vs. SVL 33.0–35.1 mm in males), vomer-
ine teeth developed (vs. absent), tibiotarsal articulation of
adpressed limb reaching between nostrils and eyes (vs.
beyond anterior corner of eye), iris distinctly bicolored,
golden-yellow in upper one-fourth and reddish brown in
lower three-fourths (vs. iris light brown with dark wash);
from A. marmoratus by tibiotarsal articulation of ad-
pressed limb reaching between nostrils and eyes (vs. be-
yond tip of snout), discontinuous glandular dorsolateral
fold from rear of eye to near vent (vs. distinct dorsolateral
fold absent), rictal gland absent (vs. multiple small glob-
ular rictal glands on right side, single rictal gland on left
side, just posterior to jaw); from A. afghanus by SVL 51.5
mm in female (vs. SVL 67.7–94.1 mm in females), rictal
gland absent (vs. indistinct rictal glands present on one
side of head at posterior end of jaw), absence of vocal
sacs in males (vs. males with dual gular pouches).
Amolops vitreus (Bain, Stuart & Orlov,
2006)
Common name. vitreous cascade frog, glass torrent frog
Type locality. collected on a stream bank near Nam
Khang River in hilly evergreen forest, Phou Dendin Na-
tional Biodiversity Conservation Area, Phongsaly Dis-
trict, Phongsaly Province, Laos.
Figure 6. Amolops vitreus (KIZ 050452). A Lateral view, B dorsal view, C ventral view of hand, D foot. Photos by Shao-Bing Hou.
Vertebrate Zoology 74, 2024, 343–357 353
Conservation status. IUCN: VU.
Type specimens of A. vitreus. Holotype: FMNH 258182,
based on original designation.
Specimen examined. Adult male (KIZ 050452) collected
on 01, August, 2022 by Yun-He Wu, Shao-Bin Hou, and
Zhong-Xiong Fu from Xishuangbanna National Nature
Reserve, Mengla, Yunnan Province, China (21.73742°N,
101.53935°E, elevation 899 m a.s.l.).
Chinese name. Based on the type locality, we suggest the
Chinese formal name as “丰沙里湍蛙”.
Morphological description (measurements in mm;
provided in Table 1). Morphological characters of the
specimen from China agreed well with the original de-
scription of Bain et al. (2006). Adult male with SVL 38.9
mm; head length (HL 15.4 mm, 39.6% of SVL) longer
than width (HW 13.9 mm, 35.7% of SVL); snout obtuse-
ly pointed in dorsal view, projecting beyond lower jaw,
round in prole, its length longer than horizontal diam-
eter of eye (ED/SL 67.7%); canthus rostralis rounded,
loreal region slightly concave, oblique; interorbital space
slightly larger than width of upper eyelid (UEW/IOS
92.7%) and internarial distance (INS/IOS 112.2%); tym-
panum distinct (TD 2.7 mm), rounded, more than half eye
diameter (ED 4.4 mm); vomerine teeth developed; tongue
cordiform, deeply notched posteriorly; pupil horizontal
(Fig. 6A); external subgular vocal sacs present, vocal sac
opening on floor of mouth at each corner.
Forelimbs slender; length of lower arm and hand
(LAHL 19.6 mm, 50.4% of SVL), about half SVL; rel-
ative nger lengths: I<II<IV<III; tips of all ngers ex-
panded into discs with circummarginal grooves; webbing
between ngers absent; subarticular tubercles distinct,
formula 1, 1, 2, 2; two metacarpal tubercles; velvety nup-
tial pad on rst nger (Fig. 6C).
Hindlimbs long, tibia (TIB 23.5 mm) more than half
SVL, longer thigh length (22.2 mm) and foot length
(FTL 20.3 mm); tibiotarsal articulation beyond the snout
when the leg is stretched forward; relative length of toes:
I<II<III<V<IV; heels overlapping when thighs are po-
sitioned at right angles to the body; tips of all toes ex-
panded into discs with circummarginal grooves; fully
webbing between toes; subarticular tubercles prominent
and rounded, formula 1, 1, 2, 3, 2; inner metatarsal tu-
bercle distinct and oval, outer metatarsal tubercle absent
(Fig. 6D).
Dorsal skin and ventral surfaces of head, body, limbs,
and flanks relatively smooth, with exception of small
tubercles posterior surface of thigh; supratympanic fold
absent; dorsolateral fold distinct, from posterior corner of
upper eyelid to near vent; two rictal glands present; hu-
meral gland absent (Fig. 6).
Figure 7. Amolops vitreus (KIZ 050452) in preservative. A Dorsal view, B ventral view, C ventral view of hand, D ventral view of
foot. Photos by Zhong-Bin Yu.
Wu Y-H et al.: Two species of the genus Amolops (Anura: Ranidae) from Yunnan border
354
Color in in life. Dorsal surface brown, with some dark
brown spots; upper lip stripe white, extending from tip
of snout to posterior of arm insertion; narrow, reddish
brown stripe on edge of canthus from tip of snout among
margin of upper eyelid, continuing along upper edge of
dorsolateral fold; dorsal surface of limbs light brown with
dark brown crossbars, interspersed with small dark brown
spots; tympanic region dark brown; throat, chest and ante-
rior part of belly light cream; flank dark brown upper one-
third, green lower two-third; expanded nger tips reddish,
except for rst nger tips yellow; subarticular tubercles
on toes, expanded toe tips, and inner metatarsal tubercle
dark brown; iris distinctly bicolored, silvery-white in up-
per one-fourth and reddish brown in lower three-fourths,
black reticulations throughout (Fig. 6).
Color in preservative. After one year of storage in etha-
nol, dorsal surface fading to grayish brown; black cross-
bars present on dorsal surfaces of limbs, ngers and toes
becoming indistinct; dorsolateral fold yellowish; throat,
chest, and abdomen cream-white; ventral surface of limbs
light yellow; ventral surface of the hands cream-white;
digit tips, subarticular tubercles of ngers, metacarpal
tubercles, and nuptial pad fading to cream-yellow or
grayish-white; toe webbing greyish brown with dark gray
flecking; ventral surface of the toes greyish brown, digit
tips, subarticular tubercles of toes and inner metatarsal
tubercle fading to greyish brown (Fig. 7).
Ecological notes. Amolops vitreus was found in rocky,
fast-flowing streams at night (20:30–23:30 h) on 01 Au-
gust 2022, surrounded by evergreen broad-leaved forest.
Other frog species observed along the stream included
Leptobrachella eos, Polypedates megacephalus, and Xe-
nophrys sp.
Distribution. This study further extends the geographical
range of A. vitreus to approximately 82.6 km from the
nearest known locality of the species. Amolops vitreus is
currently known in Phou Dendin National Biodiversity
Conservation Area, Phongsaly Province, Laos; Muong
Nhe Nature Reserve, Dien Bien Province, northwestern
Vietnam; and Xishuangbanna National Nature Reserve,
Mengla, Yunnan province, China.
Discussion
Currently, there are 22 recognized species of Amolops
known to exist in Yunnan Province (AmphibiaChina
2023). The new species and new species record described
in this study brings the total number of Amolops species
from Yunnan to 24. Our study provides more evidence
that the amphibian diversity has been largely underesti-
mated along the border regions of southern China (e.g.,
Yuan et al. 2019; Chen et al. 2020; Wu et al. 2020, 2021,
2023a,b; Zhang et al. 2022). It further unveils the taxon-
omy of some species of the genus Amolops such as A.
mengyangensis which have long been controversial (Wu
et al. 2020). Future intensied surveys and international
collaboration along these international borders combined
with molecular and acoustic data, will likely uncover
more new amphibia species and new country records.
In addition, integrating a more detailed morphological
investigation as well as assimilation of multi-loci nucle-
ar genomic markers to resolve taxonomic disputes will
speed up species discovery. This study increases the to-
tal number of Amolops species to 85, and the number of
Amolops species known from China to 53.
The Gaoligong Mountains consist of a long, narrow,
mountain range running from north to south in the west-
ern part of China’s Yunnan Province adjoining northern
Myanmar. It spans a 5° latitude with a large elevation
range of 210 m to 5000 m (Chaplin 2005). It has been
shown to have an isolating eect on many Amolops spe-
cies. For instance, A. kaulbacki and A. putaoensis are
only known to be distributed on the western slope of the
Gaoligong Mountains based on our eld work and related
researches (Frost 2023). Currently, A. kaulbacki is dis-
tributed in Kachin, Northern Myanmar and Pianma, Yun-
nan, China, and A. putaoensis in Kachin State, Myanmar
and upper Dulong River, Yunnan, China. In addition, Wu
et al. (2020) indicated that A. bellulus was paraphyletic,
forming two distinct monophyletic clusters, one of which
is distributed on the western slope of Gaoligong Moun-
tains, and the other clade is distributed on the eastern
slope of Gaoligong Mountains. Since the type locality of
A. bellulus is on the western slope (Liu et al. 2000), the
clade from the eastern slope is speculated to be a potential
new species. A similar situation was also found in A. yan-
gi sp. nov. and A. longimanus. Amolops yangi is distrib-
uted on the western slope of Gaoligong Mountains, while
A. longimanus is distributed on the eastern slope of Gaol-
igong Mountains. The Gaoligong Mountains present a
large climate dierence in the eastern slope and the west-
ern slope due to the influence of the Indian Ocean mon-
soon climate, and the rainfall on the western slope which
is signicantly greater than that on the eastern slope (Li
and Li 2020). Therefore, this distribution pattern may be
due to the unique topographic and climatic characteristics
of the Gaoligong Mountains. Future studies on compara-
tive phylogeography of these species can be carried out to
explore the formation mechanism.
Acknowledgments
This work was supported by the National Key R&D Program of Chi-
na (2022YFC2602500), the Second Tibetan Plateau Scientic Expe-
dition and Research (STEP) program (Grant No. 2019QZKK0501),
Ministry of Science and Technology of the People’s Republic of
China (2021FY100203); National Natural Science Foundation of
China (NSFC 32100371); Major Science and Technique Program
(202102AA310055), Key R & D program (202103AC100003), Yun-
nan Applied Basic Research Projects (No. 202301AT070312), Talent
and Platform of Science and Technology in Yunnan Province Science
and Technology Department (202205AM070007), Yunnan Applied Ba-
sic Research Projects (No. 202301AT070312), and the Digitalization,
Vertebrate Zoology 74, 2024, 343–357 355
Development and Application of Biotic Resource (202002AA100007)
in Yunnan Province; China’s Biodiversity Observation Network (Si-
no-BON), and the Animal Branch of the Germplasm Bank of Wild
Species, CAS (Large Research Infrastructure Funding). We thank Shao-
Bing Hou Dong An, and Tian-En Chen for their help in the eld. We
thank the Gaoligongshan National Nature Reserve and Xishuangbanna
National Nature Reserve, Yunnan, for their support to undertake eld
surveys and specimen collections.
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Supplementary Material 1
Tables S1–S5
Authors: Wu Y-H, Yu Z-B, Lu C-Q, Zhang Y-P, Dong W-J, Liu X-L, Kilunda FK, Xiong Y, Jiang Y-F, Ouyang H, Fu
Z-X, He Y-B, Yuan Z-Y, Che J (2024)
Data type: .zip
Explanation notes: Table S1. Localities, voucher information, and Genbank accession numbers for all specimens
used in this study. — Table S2. Best-tting models and partitions selected by JMODELTEST v2.1.7 for phylogeny
analysis. — Table S3. Morphological data from Amolops yangi sp. nov. and other species of A. viridimaculatus
group used in the PCA analyses. — Table S4. Uncorrected p-distance (percentage) of Amolops species included
in phylogenetic analyses and standard error estimates. — Table S5. Summary statistics and principal component
analysis scores for the mensural characters of Amolops yangi sp. nov. and other species of A. viridimaculatus group.
Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/
licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely
share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source
and author(s) are credited.
Link: https://doi.org/10.3897/vz.74.e108013.suppl1
Available via license: CC BY 4.0
Content may be subject to copyright.
