A New Species of Scutiger from Sichuan, China, Previously Misreported as S . chintingensis (Anura: Megophryidae)

Full text

Asian Herpetological Research 2025, 16(1): 20–35
DOI: 10.3724/ahr.2095-0357.2024.0049
CSTR: 32242.14.ahr.2095-0357.2024.0049
A New Species of Scutiger from Sichuan, China, Previously Misreported as
S.chintingensis (Anura: Megophryidae)
Zhitong LYU
1
, Zongyuan GAO
1,2
, Junjie HUANG
1
, Dihao WU
1
, Dechun JIANG
1,2
, Ke JIANG
1*
and Jiatang LI
1,2*
1
CAS Key Laboratory of Mountain Ecological Restoration and Bioresource Utilization, Ecological Restoration and Biodiversity Conservation
Key Laboratory of Sichuan Province, Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610213, Sichuan, China
2
University of Chinese Academy of Sciences, Beijing 100049, China
Abstract Comprehensive surveys on the distribution
of the endangered Chinting alpine toad (Scutiger
chintingensis) were recently conducted, revealing
important findings regarding its population
characteristics. This study integrated morphological
and molecular analyses to clarify the taxonomic status
of these populations. Based on detailed morphological
comparisons and phylogenetic analyses, the
population from Wolong, Wenchuan County,
Sichuan, China, is identified as a new species, which
is formally described herein. The new species can be
distinguished from all known congeners by a unique
combination of following characteristics: (1) small
body size, SVL 42.2–46.9 mm in adult males, SVL
54.3–56.3 in adult females; (2) vomerine ridges and
vomerine teeth absent, maxillary teeth present; (3)
internasal distance and interorbital distance relatively
large; (4) subarticular tubercle absent on fingers and
toes; (5) dorsal and lateral skin rough with densely-
distributed horny spines in males, smooth and lacking
black horny spines in females; (6) vocal sac absent in
males; (7) pair of pectoral glands and pair of axillary
glands present on chest in males, axillary gland about
two-thirds pectoral gland in size; (8) in breeding
males, pectoral glands and axillary glands covered by
dense spines, belly rough with horny spines, nuptial
spines present on dorsal and lateral surface of first
and second fingers and inner side of third finger,
dense spines on inner forelimb aligned, extending
from wrist to near axilla. This study highlights the
critical importance of accurate taxonomic
identification, particularly for endangered species, as
misidentifications can misguide subsequent biological
research and conservation efforts.
Keywords conservation, diversity, phylogeny, Scutiger wolong
sp. nov., taxonomy
1. Introduction
The Chinting alpine toad, Scutiger chintingensis Liu & Hu, 1960,
is a rare amphibian species endemic to Sichuan Province,
China (Fei and Ye, 2016), inhabiting the high-altitude
mountains of the western margin of the Sichuan Basin. The
species was originally discovered by Prof. Chengchao LIU in
1938 at Chinting (= Jinding), Mt. Omei (= Mt. Emei, Emeishan
City), but was initially identified as Megophrys boulengeri
(Bedriaga, 1898) (now Scutiger boulengeri) (Liu, 1950).
Subsequently, Liu and Hu (1960) described it as a new
species, S.chintingensis, based on morphological differences,
naming it after its only known locality at the time. Later
studies by Yu et al. (1983) and Inger et al. (1990) reported
additional populations from Wolong, Wenchuan County and
Mt. Wa Shan (= Mt. Wawu, Hongya County). However, for
over two decades, no new confirmed localities have been
documented (Fei et al., 2009, 2012; Fei and Ye, 2016).
Due to its restricted distribution and small population size,
S.chintingensis is listed as a Second-class National Key
ORIGINAL
ARTICLE
* Corresponding authors: Prof. Jiatang LI, from Chengdu Institute of Biology
(CIB), Chinese Academy of Sciences (CAS), Chengdu, Sichuan, China, with
his research focusing on taxonomy, phylogenetics, biogeography, genomics,
and evolution of amphibians and reptiles; Mr. Ke JIANG, from CIB, CAS,
Chengdu, Sichuan, China, with his research focusing on amphibian and
reptile taxonomy.
E-mail: lijt@cib.ac.cn (Jiatang LI); jiangke@cib.ac.cn (Ke JIANG)
Received: 17 October 2024 Accepted: 27 November 2024
Published Online: 13 December 2024
2095-0357/© 2025 Asian Herpetological Research Editorial Office and
Science Press. This is an open access article under the CC BY-NC-ND
license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Protected Wild Animals of China (National Forestry and
Grassland Administration of China, 2021) and categorized as
Endangered (EN) in the China Red List of Biodiversity (Jiang
et al., 2021). Despite these protective measures and heightened
attention, critical ecological and biological information about
this species remains scarce. Notably, no substantial
population-based studies have been conducted since Xie et
al. (2000), who investigated its population ecology at Mt.
Wawu.
To address this knowledge gap, field surveys have been
conducted since 2018 by multiple teams from the Chengdu
Institute of Biology, Chinese Academy of Sciences, across the
potential distribution range of S.chintingensis. These efforts led
to the identification of a fourth confirmed locality in
Daxiangling Nature Reserve, Yingjing County, as well as
updated assessments of population sizes and threat factors for
the populations at Mt. Emei and Mt. Wawu (Fan et al., 2024).
This study focuses on the taxonomic status of S.chintingensis
through comprehensive morphological and molecular
analyses of populations from Mt. Emei, Mt. Wawu, and
Wolong. Morphological comparisons and phylogenetic
analyses revealed that the Wolong population exhibits
significant divergence from those at Mt. Emei and Mt.
Wawu, warranting its recognition as a new species. This
taxonomic revision refines the understanding of S.chintingensis
as a species with a more restricted distribution and smaller
population than previously recognized. These findings
underscore the critical role of taxonomy in conservation
biology, as misidentification of endangered species can
compromise ecological research and hinder the development
of effective conservation strategies.
2. Materials and Methods
2.1. Morphology A total of 28 specimens were examined
and measured, including the holotype of S.chintingensis, newly
collected specimens from Mt. Emei, Emeishan City, and Mt.
Wawu, Hongya County, and eight Scutiger Theobald, 1868
specimens from Wolong, Wenchuan County. All specimens
were deposited in the Herpetological Museum, Chengdu
Institute of Biology (CIB), Chinese Academy of Sciences.
External measurements followed Jiang and Li (2021) and
included snout–vent length (SVL), head length (HL), head
width (HW), snout length (SL), internasal distance (IND),
interorbital distance (IOD), horizontal diameter of eye (ED),
hand length (HAL), forearm length (FOL), length of tarsus and
foot (TFL), and tibial length (TB). All measurements were
made with digital calipers to the nearest 0.1 mm.
Statistical analyses of morphometric measurements were
conducted using R v4.0.0, following the methods described in
Lyu et al. (2023a). Analyses included one-way analysis of
variance (ANOVA) and principal component analysis (PCA),
with all measurements logarithmically transformed prior to
analysis. Only male specimens (seven from Mt. Emei
population, ten from Mt. Wawu population, and six from
Wolong population) were included in analysis due to the
limited availability of female specimens (three from Mt. Emei,
none from Mt. Wawu, and two from Wolong).
Morphological comparisons with all known Scutiger
congeners were based on data from the original and
supplemental descriptions in the literature (Blyth, 1855;
Günther, 1896; Bedriaga, 1898; Liu, 1950; Liu and Hu,
1960; Dubois, 1974, 1978, 1979; Fei, 1977; Liu et al., 1978,
1979; Yang et al., 1979; Fang, 1985; Huang, 1985; Fei and Ye,
1986, 2016; Fei et al., 1995, 2009, 2012; Delorme and Dubois,
2001; Ye and Fei, 2007; Jiang et al., 2012, 2016; Khatiwada et
al., 2019; Yang and Huang, 2019; Rao, 2022 “2020”; Shi et al.,
2023; Zhou et al., 2023; Hofmann et al., 2024).
2.2. Phylogeny Liver tissue samples were collected from 16
newly obtained Scutiger specimens, including four from Mt.
Emei, four from Mt. Wawu, and eight from Wolong. Tissues
were extracted from previously anesthetized and euthanized
specimens. The samples were preserved in 95% ethanol and
stored at –40 °C. Genomic DNA was isolated using a DNA
extraction kit (Sangon Biotech Co., Ltd., Shanghai, China).
Phylogenetic analyses targeted two mitochondrional genes,
partial cytochrome C oxidase 1 (COI) and partial cytochrome
b (Cyt b), which were amplified with primers following
previously published protocols (Jia et al., 2024). Polymerase
chain reaction (PCR) amplifications were carried out under
the following cycling conditions: initial denaturation at 95 °C
for 4 min, 35 cycles of denaturation at 95 °C for 40 s,
annealing at 48 °C for 40 s, and extension at 72 °C for 60 s,
with a final extension step at 72 °C for 10 min. The PCR
products were purified using spin columns and sequenced
bidirectionally with both forward and reverse primers
(Sangon Biotech Co., Ltd., Shanghai, China).
All newly generated sequences were deposited in GenBank
(Table 1). For phylogenetic analyses, the dataset was
augmented with sequences from additional Scutiger
congeners retrieved from GenBank (Table 1), with two S.
wuguanfui Jiang, Rao, Yuan, Wang, Li, Hou, Che & Che, 2012
individuals used as the out-group based on prior phylogenetic
studies. DNA sequences were aligned using the Clustal W
algorithm (Thompson et al., 1997). PartitionFinder2 was
employed to determine the optimal partitioning scheme, while
jModelTest v2.1.2 was used to identify the best-fitting
nucleotide substitution model. Phylogenetic analyses were
conducted using maximum-likelihood (ML) in RaxmlGUI
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Table 1 Localities, voucher information, and GenBank accession numbers for all samples used in this study.
ID Species Voucher Locality COI Cyt b
1Scutiger wolong sp. nov. CIB 121679 Wolong, Wenchuan, Sichuan, China PQ585688 PQ587079
2Scutiger wolong sp. nov. CIB 121680 Wolong, Wenchuan, Sichuan, China PQ585689 PQ587080
3Scutiger wolong sp. nov. CIB 121681 Wolong, Wenchuan, Sichuan, China PQ585690 PQ587081
4Scutiger wolong sp. nov. CIB 121682 Wolong, Wenchuan, Sichuan, China PQ585691 PQ587082
5Scutiger wolong sp. nov. CIB 121683 Wolong, Wenchuan, Sichuan, China PQ585692 PQ587083
6Scutiger wolong sp. nov. CIB 121684 Wolong, Wenchuan, Sichuan, China PQ585693 PQ587084
7Scutiger wolong sp. nov. CIB 121685 Wolong, Wenchuan, Sichuan, China PQ585694 /
8Scutiger wolong sp. nov. CIB 121686 Wolong, Wenchuan, Sichuan, China PQ585695 PQ587085
9Scutiger chintingensis CIB 121687 Jinding, Mt. Emei, Emeishan, Sichuan, China PQ585696 PQ587086
10 Scutiger chintingensis CIB 121689 Jinding, Mt. Emei, Emeishan, Sichuan, China PQ585697 PQ587087
11 Scutiger chintingensis CIB 121690 Jinding, Mt. Emei, Emeishan, Sichuan, China PQ585698 PQ587088
12 Scutiger chintingensis CIB 121691 Jinding, Mt. Emei, Emeishan, Sichuan, China PQ585699 PQ587089
13 Scutiger chintingensis CIB 121695 Mt. Wawu, Hongya, Sichuan, China PQ585700 PQ587090
14 Scutiger chintingensis CIB 121698 Mt. Wawu, Hongya, Sichuan, China PQ585701 PQ587091
15 Scutiger chintingensis CIB 121699 Mt. Wawu, Hongya, Sichuan, China PQ585702 PQ587092
16 Scutiger chintingensis CIB 121700 Mt. Wawu, Hongya, Sichuan, China PQ585703 PQ587093
17 Scutiger feiliangi SYAU BAA000040 Baiyunshan, Songxian, Henan, China OR263444 OR257694
18 Scutiger feiliangi bys1 Baiyunshan, Songxian, Henan, China KF757397 KF757340
19 Scutiger feiliangi ljs18 Laojunshan, Luanchuan, Henan, China KF757425 KF757353
20 Scutiger feiliangi srs15 Shirenshan, Lushan, Henan, China KF757437 KF757362
21 Scutiger feiliangi srs9 Shirenshan, Lushan, Henan, China / KF757391
22 Scutiger ningshanensis nsc1 Pingheliang, Ningshan, Shaanxi, China KF757427 KF757355
23 Scutiger ningshanensis nsc11 Pingheliang, Ningshan, Shaanxi, China KF757429 KF757383
24 Scutiger ningshanensis nsc2 Pingheliang, Ningshan, Shaanxi, China KF757433 KF757386
25 Scutiger ningshanensis nsc9 Pingheliang, Ningshan, Shaanxi, China KF757436 /
26 Scutiger boulengeri A1-AL Tagejia, Ngamring, Xizang, China KY310870 KY310922
27 Scutiger boulengeri JS1507_C1 Lhasa, Xizang, China KY310875 KY310927
28 Scutiger ghunsa JRK2015-193 Ghunsa, Taplejung, Nepal MK970591 MK970614
29 Scutiger ghunsa JRK2015-205 Ghunsa, Taplejung, Nepal MK970603 MK970626
30 Scutiger glandulatus SC1_2014 Garze, Sichuan, China KY310879 KY310934
31 Scutiger glandulatus CIB XM1188 Xiangcheng, Sichuan, China / FJ945466
32 Scutiger gongshanensis KIZ 020492 Yunnan, China MW021395 MW133458
33 Scutiger jiulongensis KIZ 045055 Garze, Sichuan, China KU243066 MW133417
34 Scutiger jiulongensis KIZ YPX32464 Sichuan, China MW021354 MW133416
35 Scutiger kanjiroba NHME A0576/99 Dolpa, Nepal KY310896 KY310947
36 Scutiger kanjiroba A11_12 Baglung, Nepal KY310890 KY310941
37 Scutiger liupanensis FWR11 Fragrant Water River, Jingyuan, Ningxia, China KC140483 JX533792
38 Scutiger liupanensis DDR6 Double-Dragon River, Jingyuan, Ningxia, China KC140551 JX533734
39 Scutiger luozhaensis CIB 119115 Gari, Lhozhag, Xizang, China OR141832 /
40 Scutiger luozhaensis CIB 119119 Gari, Lhozhag, Xizang, China OR141823 /
41 Scutiger mammatus CIB XM949 Jiagenba, Kangding, Sichuan, China MW021360 FJ463152
42 Scutiger muliensis KIZ YPX36717 Sichuan, China MW167046 MW133420
43 Scutiger muliensis KIZ YPX36718 Sichuan, China MW167047 MW133421
44 Scutiger nepalensis A2018/13 Tribeni, Khaptad, Chainpur, Nepal KY310886 KY310937
45 Scutiger nyingchiensis BQ16 Lulang, Bayi, Xizang, China OM506176 OM505290
46 Scutiger nyingchiensis lz10 Lulang, Bayi, Xizang, China OM506169 OM505283
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v1.3 (Silvestro and Michalak, 2012) and Bayesian inference
(BI) in MrBayes v3.2.4 (Ronquist et al., 2012). For ML analysis,
evolutionary relationships of the analyzed taxa were inferred
from a bootstrap consensus tree generated from 1 000
replicates. For BI analysis, two independent runs were
conducted, each spanning 10 000 000 generations with
sampling every 1 000 generations. The first 25% of samples
were discarded as burn-in, resulting in a potential scale
reduction factor (PSRF) of <0.005. Genetic distances between
and within species were calculated in MEGA 6 using the
uncorrected p-distance model, with pairwise deletion of gaps
and missing data.
3. Results
The ML and BI analyses yielded congruent topologies. As a
result, the ML phylogeny is presented, with bootstrap support
(BS) values and Bayesian posterior probabilities (BPP) labeled
at the nodes (Figure 1). Phylogenetic analyses revealed that the
Scutiger populations from the type locality at Mt. Emei (IDs 9–
12) and the neighboring Mt. Wawu (IDs 13–16) formed a
strongly supported clade (BS=100, BPP=1.00), with minimal
genetic divergence (p-distance 0.0–0.7% in COI and 0.0–1.0%
in Cyt b), representing the S.chintingensis lineage. In contrast,
the Scutiger population from Wolong (IDs 1–8), previously
recorded as S.chintingensis, formed a distinct lineage with
robust support (BS=100, BPP=1.00) and negligible genetic
divergence (p-distance 0.0–0.7% in COI and 0.0–0.6% in Cyt b).
The Wolong lineage was recovered as the sister taxon to the S.
chintingensis lineage (BS=100, BPP=1.00), displaying clear
interspecific genetic divergence (p-distance 5.7%–6.9% in
COI and 5.2%–6.2% in Cyt b), which is significant within the
genus (Table 2). Together, these two lineages formed a sister
group (BS=92, BPP=1.00) to the clade comprising S.feiliangi
Zhou, Guan & Shi, 2023 and S.ningshanensis Fang, 1985.
Morphological analyses further supported these findings.
Statistical evaluations of morphometric measurements
indicated that specimens from Mt. Emei and Mt. Wawu
were morphologically identical, whereas those from Wolong
exhibited significant differences. PCA showed that the first
two principal components (PCs) accounted for 56.5% and
16.9% of the variance, respectively, explaining over 70.0% of
the total variance. Scatter plots demonstrated clustering of S.
chintingensis specimens from Mt. Emei and Mt. Wawu, while
specimens from Wolong formed a separate, isolated cluster
(Figure 2A). ANOVA revealed significant differences in HL,
IOD, IND, and TB between the Wolong population and S.
chintingensis (P<0.05; Table 3; Figure 2B–L). Additionally,
specimens from Wolong exhibited a distinct combination of
morphological characteristics, not only distinguishing them
from S.chintingensis populations at Mt. Emei and Mt. Wawu
but also from all other congeners within the genus (presented
in detail below).
Taken together, the morphological and phylogenetic
evidence unequivocally supports the recognition of the
Wolong population as a new species, which is described
below.
4. Taxonomic account
4.1. Scutiger wolong sp. nov.
Chresonymy Scutiger chintingensis—Yu et al., 1983; Fan et al.,
2024 (Wolong)
Scutiger (Scutiger)chintingensis—Fei et al., 2009 (Wenchuan);
Fei et al., 2012 (Wenchuan); Fei and Ye, 2016 (Wenchuan)
Holotype CIB 121683 (Figure 3), adult male, collected by
Continued Table 1
ID Species Voucher Locality COI Cyt b
47 Scutiger occidentalis Pk6 Deosai, Pakistan KY310901 KY310957
48 Scutiger occidentalis Pk5 Deosai, Pakistan KY310900 KY310956
49 Scutiger sikimmensis KIZ 011127 Yadong, Xizang, China KU243058 MW133429
50 Scutiger sikimmensis KIZ 013982 Yadong, Xizang, China MW021367 MW133431
51 Scutiger spinosus KIZ 011100 Medog, Xizang, China KU243054 MW133427
52 Scutiger spinosus KIZ 011092 Medog, Xizang, China MW021362 MW133426
53 Scutiger tengchongensis SYS a005799 Tengchong, Yunnan, China MK121783 /
54 Scutiger tengchongensis SYS a005800 Tengchong, Yunnan, China MK121784 /
55 Scutiger tuberculatus CIB XM995 Laji, Yuexi, Sichuan, China MW021351 FJ945492
56 Scutiger tuberculatus KIZ YPX11046 Sichuan, China MW021397 MW133413
57 Scutiger wanglangensis 21514N1 Beichuan, Sichuan, China OQ361635 /
58 Scutiger wanglangensis 21505N4 Beichuan, Sichuan, China OQ361637 /
59 Scutiger wuguanfui KIZ 011101 Medog, Xizang, China KU243060 MW133433
60 Scutiger wuguanfui KIZ 011102 Medog, Xizang, China KU243061 MW133434
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23

Junjie HUANG on 31 May 2020 from Wolong Town
(30.9198° N, 103.0580° E, ca 2 414 m a.s.l.), Wenchuan
County, Sichuan Province, China.
Paratypes CIB 121679–121682, 121684, five adult males,
and CIB 121685–121686, two adult females, collected at the
same time as the holotype.
Etymology The specific name wolong is used as a noun in
apposition and refers to Wolong Town, the type locality of this species.
Common name Wolong alpine toad (in English)/wò lóng
chǐ tū chán (卧龙齿突蟾 in Chinese).
Diagnosis Scutiger wolong sp. nov. can be diagnosed by the
following combination of characteristics: (1) small body size,
SVL 42.2–46.9 (44.8±2.1, n=6) mm in adult males, SVL 54.3–
56.3 mm (n=2) in adult females; (2) vomerine ridges and
vomerine teeth absent, maxillary teeth present; (3) internasal
distance and interorbital distance relatively large; (4)
0.02
14
38
36
21
35
31
3
48
20
52
23
9
57
12
29
55
40
45
11
15
27
30
59
22
46
56
17
47
7
60
4
44
1
49
28
53
51
5
32
58
41
13
50
39
10
2
8
16
43
24
25
6
34
33
19
37
18
42
26
54
wuguanfui
boulengeri
glandulatus
mammatus
muliensis
tuberculatus
jiulongensis
liupanensis
wanglangensis
occidentalis
spinosus
tengchongensis
gongshanensis
luozhaensis
nyingchiensis
ghunsa
nepalensis
kanjiroba
sikimmensis
ningshanensis
feiliangi
chintingensis
wolong sp. nov.
Scutiger
100/1.00
100/1.00
100/1.00
96/1.00
97/1.00
100/1.00
92/1.00
100/1.00
100/1.00
100/1.00
100/1.00
70/0.98
100/1.00
96/1.00
93/1.00
60/0.85
100/1.00
87/1.00
100/1.00
94/1.00
100/1.00
97/1.00
99/1.00
99/1.00
99/1.00
100/1.00
100/1.00
100/1.00
100/1.00
100/1.00
100/1.00
100/1.00
77/1.00
100/1.00
100/1.00
BS/BPP
Figure 1 Maximum-likelihood phylogeny for Scutiger based on mitochondrial COI-Cyt b gene fragments. Bootstrap support (BS)>60 and
Bayesian posterior probability (BPP)>0.85 are shown. Number at ends of lineages correspond to IDs in Table 1.
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Table 2 Range of p-distance (in 0.1%) based on COI and CYTB genes among all Scutiger species used in this study.
ID Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
Based on COI gene
1S.wolong sp.
nov. 0.0–
0.7
2S.chintingensis 5.7–
6.9 0.0–
0.7
3S.feiliangi 10.7–
11.4 11.1–
11.6 0.0–
0.3
4S.ningshanensis 10.0–
10.7 11.3–
11.8 1.0–
1.7 0.0–
0.5
5S.boulengeri 12.1–
12.6 10.8–
12.1 11.1–
12.0 11.5–
12.4 0.5
6S.ghunsa 10.6–
10.8 9.6–
10.3 11.7–
12.2 11.6–
12.1 10.3–
10.5 0.0
7S.glandulatus 11.8–
12.2 11.7–
12.6 11.8–
12.1 11.8–
12.3 6.3–
6.3 11.1–
11.1 /
8S.gongshanensis 11.3–
11.8 10.1–
10.7 10.6–
10.9 10.4–
10.8 11.0–
11.2 8.2–
8.2 11.3 /
9S.jiulongensis 11.8–
12.8 10.2–
11.8 10.7–
11.4 10.9–
11.8 5.3–
5.7 10.6–
11.5 5.9–
5.9 10.2–
11.1 1.2
10 S.kanjiroba 11.7–
13.6 11.2–
11.8 9.9–
10.6 10.8–
11.5 10.7–
11.6 7.5–
7.9 10.6–
11.0 9.0–
9.4 10.6–
11.4 1.6
11 S.liupanensis 11.0–
12.7 9.7–
10.2 10.2–
11.6 10.6–
12.2 5.8–
6.1 9.7–
10.0 5.3–
5.7 9.3–
10.2 3.6–
4.0 9.5–
9.9 2.1
12 S.luozhaensis 12.9–
13.7 11.7–
12.1 10.4–
10.6 10.2–
10.6 12.6–
12.9 10.1–
10.1 12.5–
12.5 7.5–
7.5 11.8–
12.3 10.5–
11.4 11.8–
12.1 0.0
13 S.mammatus 11.9–
12.4 11.6–
12.5 11.3–
11.5 11.2–
11.7 6.2–
6.4 10.7–
10.7 6.9 10.1 5.1–
5.1 9.7–
10.1 4.9–
5.7 12.4–
12.4 /
14 S.muliensis 11.2–
12.2 9.9–
11.2 9.9–
10.6 10.3–
11.0 5.7–
6.4 11.4–
11.6 6.7–
6.9 11.7–
11.9 5.3–
5.9 9.2–
10.3 5.7–
5.9 11.7–
11.9 5.7–
11.9 0.5
15 S.nepalensis 12.1–
12.9 11.2–
11.4 11.7–
12.2 12.4–
12.8 9.6–
9.9 7.9–
7.9 10.4 8.8 9.3–
9.3 5.6–
5.8 8.2–
8.9 11.9–
11.9 10.1 9.4–
9.6 /
16 S.nyingchiensis 11.5–
12.2 10.6–
11.6 9.1–
10.7 9.3–
10.8 10.8–
11.3 9.0–
9.4 11.6–
11.8 5.9–
6.1 10.8–
12.1 9.9–
11.0 10.1–
11.1 5.9–
6.7 10.7–
11.5 11.5–
12.9 10.4–
11.0 3.0
17 S.occidentalis 10.6–
10.8 10.3–
11.2 10.6–
10.9 10.2–
10.8 9.9–
10.8 7.5–
8.9 9.9–
10.4 8.6–
9.9 8.7–
9.6 8.6–
9.6 9.3–
10.7 10.6–
10.8 9.2–
9.3 8.2–
9.7 8.8–
8.8 9.5–
10.4 1.4
18 S.sikimmensis 12.3–
13.0 11.2–
12.1 10.2–
10.9 9.3–
10.0 12.6–
13.7 10.5–
11.0 12.7–
13.2 11.4–
11.4 13.1–
13.5 11.6–
12.5 11.7–
13.1 10.5–
11.0 12.4–
12.8 12.6–
13.3 12.5–
13.0 9.9–
10.7 11.2–
11.7 0.3
No. 1
Zhitong LYU et al.
A New Species of Scutiger
25

Continued Table 2
ID Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
19 S.spinosus 10.0–
10.7 9.6–
9.8 9.9–
10.1 9.6–
10.1 10.7–
10.9 8.1–
8.1 10.4–
10.4 6.3–
6.3 11.2–
11.7 7.1–
7.5 9.9–
10.4 8.7–
8.7 11.4–
11.4 10.1–
10.3 8.5–
8.5 6.7–
7.7 9.4–
10.5 10.3–
10.7 0.0
20 S.tengchongensis 11.5–
12.2 10.5–
11.0 10.6–
11.1 10.6–
11.1 9.9–
10.4 10.3–
10.5 10.5–
10.7 6.9–
7.1 10.0–
11.1 9.2–
9.9 9.1–
10.2 9.0–
9.2 10.8–
11.0 11.5–
12.0 9.9–
10.1 8.1–
8.7 10.3–
10.6 11.6–
12.3 7.5–
7.7 0.2
21 S.tuberculatus 11.3–
11.8 10.6–
11.0 10.4–
10.6 9.9–
10.4 6.6–
7.2 12.2–
12.2 7.1–
7.1 11.5–
11.5 6.5–
7.3 11.2–
11.4 5.9–
6.4 12.3–
12.3 6.3–
6.3 4.1–
4.3 10.6–
10.6 11.9–
12.7 8.6–
9.9 11.9–
12.4 10.3–
10.3 10.6–
10.8 0.0
22 S.wanglangensis 12.0–
13.4 10.6–
11.7 9.8–
10.5 10.6–
11.5 6.4–
6.9 10.8–
11.3 6.5–
6.7 10.1–
10.6 4.3–
4.5 9.3–
10.1 1.6–
3.2 11.8–
12.3 4.9–
5.1 5.1–
5.5 9.3–
9.3 11.1–
11.8 9.9–
10.4 12.2–
13.1 10.1–
10.6 10.4–
10.7 5.8–
5.8 0.5
23 S.wuguanfui 13.6–
14.2 12.2–
12.4 11.5–
11.7 11.5–
11.7 11.9–
12.4 12.6–
12.6 13.6–
13.6 12.2–
12.2 13.4–
13.8 13.5–
13.8 12–
13.3 14.3–
14.3 12.6–
12.6 12.8–
13.0 11.7–
11.7 11.7–
12.3 12.8–
12.8 11.3–
11.8 12.0–
12.0 12.6–
12.9 12.0–
12.0 12.1–
12.6 0.0
Based on Cyt b gene
1S.wolong sp.
nov. 0.0–
0.6
2S.chintingensis 5.2–
6.2 0.0–
1.0
3S.feiliangi 11.3–
12.7 11.8–
13.5 0.0–
1.0
4S.ningshanensis 10.6–
12.4 11.5–
13.2 2.0–
3.0 0.0–
1.4
5S.boulengeri 13.4–
14.2 12.5–
13.7 13.5–
14.7 14.6–
21.1 0.4
6S.ghunsa 10.8–
11.1 11.1–
11.8 13.7–
14.8 12.7–
14.4 12.9–
12.9 0.0
7S.glandulatus 13.4–
14.2 12.9–
14.4 15.2–
16.2 15.4–
16.4 7.1–
8.0 12.7–
13.2 0.4
8S.gongshanensis 15.6–
15.8 16.3–
17.4 18.3–
19.4 17.5–
18.3 15.5–
16.0 15.9–
15.9 14.2–
14.5 /
9S.jiulongensis 13.0–
13.5 13.7–
14.7 15.0–
16.3 14.2–
16.2 9.8–
10.5 11.5–
11.7 8.8–
9.3 15.7–
16.0 0.2
10 S.kanjiroba 13.5–
14.5 13.0–
15.2 15.2–
16.0 14.9–
16.7 13.0–
14.2 13.2–
13.2 13.4–
13.9 15.4–
16.4 11.3–
12.0 4.1
11 S.liupanensis 12.7–
13.7 12.2–
14.0 15.8–
17.1 15.4–
17.5 7.3–
8.5 10.8–
12.0 4.9–
5.8 13.7–
13.7 6.2–
7.7 12.5–
13.2 2.4
12 S.luozhaensis / / / / / / / / / / / /
13 S.mammatus 12.7–
13.0 12.4–
13.4 14.3–
15.3 14.2–
15.4 7.1–
7.6 11.3–
11.3 6.0–
6.4 14.5 7.7–
8.0 12.4–
12.4 5.8–
7.6 / /
14 S.muliensis 12.3–
13.2 12.8–
14.0 13.7–
14.7 14.4–
15.4 7.2–
8.3 11.6–
12.3 6.4–
6.9 12.8–
13.5 9.3–
10.2 12.5–
13.7 7.1–
8.2 /4.3–
5.0 1.0
Vol. 16
Asian
Herpetological
Research
26

subarticular tubercle absent on fingers and toes; (5) dorsal and
lateral skin rough with densely-distributed horny spines in
males, smooth and lacking black horny spines in females; (6)
vocal sac absent in males; (7) pair of pectoral glands and pair of
axillary glands present on chest in males, axillary gland about
two-thirds of pectoral gland in size; (8) in breeding males,
pectoral glands and axillary glands covered by dense spines,
belly rough with horny spines, nuptial spines present on dorsal
and lateral surface of first and second fingers and inner side of
third finger, dense spines on inner forelimb aligned, extending
from wrist to near axilla.
Comparison Scutiger wolong sp. nov., previously reported as
S.chintingensis and as its sister taxon, can be readily
distinguished based on larger internasal distance
4.2–4.9 mm (vs. smaller internasal distance 3.3–4.3 mm),
larger interorbital distance 4.3–4.8 mm (vs. smaller
interorbital distance 3.1–3.7 mm), and belly rough with
horny spines in males (vs. belly smooth without spines).
Phylogenetically, Scutiger wolong sp. nov. is closely related to
S.feiliangi and S.ningshanensis. However, the new species can
be distinguished from S.feiliangi by belly rough with horny
spines in males (vs. belly smooth without spines in males),
ventral surface without spines in females (vs. large dense
spines on chest and two sides of belly in females), and
intermittent longitudinal folds on dorsum absent (vs. four lines
of intermittent longitudinal folds present on dorsum). The
new species can also be distinguished from S.ningshanensis
based on intermittent longitudinal folds on dorsum absent (vs.
four lines of intermittent longitudinal folds present on
dorsum), inner forelimb with dense spines present in
breeding males (vs. absent), and axillary gland about two-
thirds of pectoral gland in size (vs. almost equal).
Morphologically, Fei et al. (2009) noted that S.pingwuensis
Liu & Tian, 1978 is similar to S.chintingensis due to the
presence of dense spines on the inner forelimb in breeding
males. Although molecular data for S.pingwuensis are
unavailable, Scutiger wolong sp. nov. can be distinguished
from S.pingwuensis by smaller body size (SVL 42.2–46.9 mm
vs. SVL 60.7–75.8 mm in adult males) and maxillary teeth
present (vs. absent).
Among the remaining 26 congeners, Scutiger wolong sp. nov.
can be readily distinguished seriatim by vocal sac absent in
males (vs. present in S.adungensis Dubois, 1979, S.gongshanensis
Yang & Su, 1979, and S.wuguanfui), belly rough with horny
spines in males (vs. belly smooth without spines in males in S.
bangdaensis Rao, Hui, Ma & Zhu, 2022 “2020”, S.biluoensis Rao,
Hui, Zhu & Ma, 2022 “2020”, S.brevipes (Liu, 1950), S.ghunsa
Khatiwada, Shu, Subedi, Wang, Ohler, Cannatella, Xie & Jiang,
2019, S.glandulatus (Liu, 1950), S.jiulongensis Fei, Ye & Jiang,
1995, S.kanjiroba Hofmann, Jablonski & Schmidt, 2024, S.
Continued Table 2
ID Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23
15 S.nepalensis 13.7–
14.0 13.0–
14.2 16.0–
17.0 16.2–
16.7 13.5–
13.9 12.7–
12.7 13.9–
13.9 16.3 11.8–
12.0 8.7–
9.4 13.5–
14.7 / 13.0 13.0–
13.5 /
16 S.nyingchiensis 13.3–
15.2 12.6–
15.7 15.9–
18.4 16.6–
18.0 13.4–
15.5 14.1–
15.3 12.0–
14.5 11.2–
12.4 13.9–
16.0 14.3–
16.1 13.0–
14.5 /13.7–
15.7 12.3–
14.6 13.1–
14.9 4.7
17 S.occidentalis 12.5–
13.2 11.3–
13.5 12.8–
14.3 13.0–
14.5 11.2–
12.1 10.8–
11.7 11.4–
12.4 13.0–
13.0 11.9–
13.6 12.2–
13.2 10.8–
12.0 /12.4–
12.9 11.7–
12.7 12.2–
12.7 12.5–
14.7 1.2
18 S.sikimmensis 11.6–
12.3 12.7–
14.2 14.6–
16.1 15.0–
16.5 13.7–
14.7 12.0–
12.5 13.6–
14.1 15.3–
15.3 12.9–
13.2 12.5–
13.2 12.5–
13.4 /13.9–
14.3 12.7–
13.4 12.7–
13.2 13.5–
15.3 11.7–
12.2 0.4
19 S.spinosus 11.3–
12.0 11.8–
13.0 15.5–
16.8 15.3–
16.5 11.3–
12.2 13.0–
13.5 12.7–
13.4 11.5–
11.5 13.1–
13.9 13.5–
15.2 11.1–
12.7 /13.4–
13.9 13.0–
13.7 12.8–
13.3 11.8–
12.3 9.6–
10.6 11.3–
11.8 0.4
20 S.tengchongensis / / / / / / / / / / / / / / / / / / / /
21 S.tuberculatus 12.8–
13.0 13.0–
14.0 14.0–
14.8 14.5–
15.5 7.4–
7.8 12.5–
12.5 6.9–
7.3 13.2–
13.2 9.7–
10.0 12.7–
13.2 7.3–
8.2 /5.2–
5.2 2.4–
2.6 14.0–
14.0 12.8–
14.5 12.2–
12.7 13.4–
13.9 13.4–
13.9 / 0.0
22 S.wanglangensis / / / / / / / / / / / / / / / / / / / / / /
23 S.wuguanfui 21.3–
22.1 19.8–
20.6 21.1–
21.9 20.7–
21.3 16.6–
17.1 19.9–
19.9 17.1–
17.6 17.0–
17.0 17.4–
17.7 19.2–
19.4 17.9–
18.2 /18.4–
18.4 18.6–
18.9 20.8–
20.8 17.7–
19.9 17.6–
17.8 19.9–
19.9 18.5–
19.0 /19.2–
19.2 / 0.0
No. 1
Zhitong LYU et al.
A New Species of Scutiger
27

luozhaensis Shi, Sui, Ma, Ji, Bu-Dian & Jiang, 2023, S.maculatus
(Liu, 1950), S.mammatus (Günther, 1896), S.meiliensis Rao, Hui,
Zhu & Ma, 2022 “2020”, S.muliensis Fei & Ye, 1986, S.
nepalensis Dubois, 1974, S.nyingchiensis Fei, 1977, S.occidentalis
Dubois, 1978, S.sikimmensis (Blyth, 1855), S.spinosus Jiang,
Wang, Li & Che, 2016, S.tengchongensis Yang & Huang, 2019,
and S.tuberculatus Liu & Fei, 1979), inner forelimb with dense
spines present in breeding males (vs. absent in S.boulengeri,S.
liupanensis Huang, 1985, and S.wanglangensis Ye & Fei, 2007),
and maxillary teeth present (vs. absent in S.bhutanensis
Delorme & Dubois, 2001).
Description of holotype CIB 121683, adult male, SVL
46.9 mm; head width slightly longer than head length, HW/
HL 1.03; snout rounded, sloping posteriorly to mouth in
profile, protruding well beyond margin of lower jaw; canthus
rostralis well developed; nostril ovoid; loreal region slightly
oblique and concave; eye moderate, ED/HL 0.35; pupil
vertical; dorsal surface of head flat; tympanum absent; choanae
ovoid, widely separated; vomerine ridges and vomerine teeth
absent; maxillary teeth present; margin of tongue slightly
notched; vocal sac absent.
Lower arm length 0.21 of SVL and hand length 0.26 of SVL;
relative finger lengths I<II<IV<III; tip of finger rounded,
slightly dilated; fingers without webs and lateral fringes;
subarticular tubercle absent; outer and inner metacarpal
tubercles extremely flat, almost invisible; nuptial spines on
dorsal and lateral surface of first and second fingers, and inner
side of third finger.
Shank length 0.41 of SVL and tarsus-foot 0.67 of SVL; tibio-
tarsal articulation reaching temporal region when hindlimb
stretched alongside body; heels not meeting when hindlimbs
held at right angles to body; relative toe lengths I<II<V=III<IV;
tips of toes rounded, slightly dilated; developed webs and
lateral fringes present between toes; subarticular tubercle
absent; inner metatarsal tubercle long and indistinct, outer
metatarsal tubercle absent; tarsal folds absent.
Dorsal and lateral skin very rough; densely distributed
horny spines on head, including snout, loreal and temporal
regions, lip, upper eyelid, and supratympanic fold;
supratympanic fold distinct and thick, extending from
−0.1
0.0
0.1
0.2
−0.2 −0.1 0.0 0.1
PC1 (56.5%)
PC2 (16.9%)
37.5
40.0
42.5
45.0
47.5
chintingensis wolong sp. nov.
SVL
13.5
14.0
14.5
15.0
15.5
16.0
chintingensis wolong sp. nov.
HL (P<0.05)
14
15
chintingensis wolong sp. nov.
HW
5.6
6.0
6.4
6.8
chintingensis wolong sp. nov.
SL
3.5
4.0
4.5
chintingensis wolong sp. nov.
IND (P<0.001)
3.5
4.0
4.5
chintingensis wolong sp. nov.
IOD (P<0.001)
3.5
4.0
4.5
5.0
5.5
chintingensis wolong sp. nov.
ED
10.5
11.0
11.5
12.0
12.5
13.0
chintingensis wolong sp. nov.
HAL
9.5
10.0
10.5
11.0
11.5
chintingensis wolong sp. nov.
FOL
28
30
32
chintingensis wolong sp. nov.
TFL
16
17
18
19
chintingensis wolong sp. nov.
TB (P<0.01)
chintingensis Mt. Emei
chintingensis Mt. Wawu
wolong sp. nov.
A B C D
EF G H
I J K L
Figure 2 Morphometric comparisons of male specimens of Scutiger chintingensis (n=17) and S.wolong sp. nov. (n=6). A: Scatter plot of PC1 and
PC2 from Principal Component Analysis; B–L: Boxplots for all 11 morphological variables.
Vol. 16
Asian
Herpetological
Research
28

posterior part of upper eyelids to shoulder; densely distributed
horny spines and several scattered large tubercules on dorsum;
flank and dorsolateral limbs with densely distributed horny
spines, those on lower arm and inner upper arm larger and
translucent. Ventral skin rough; horny spines densely
distributed on lower lip, rather sparse on throat; pair of
Table 3 Morphometric comparisons based on the morphometric measurements (in mm) of male specimens of Scutiger wolong sp. nov. (n=6) and S.
chintingensis (n=17). *Pvalues<0.05, **Pvalues<0.01, ***Pvalues<0.001.
S.wolong sp. nov. S.chintingensis P value
SVL 42.2–46.9 (44.8±2.1) 37.5–47.2 (43.2±2.6) 0.1907
HL 14.2–16.0 (15.1±0.6) 13.6–15.8 (14.2±0.6) 0.0280*
HW 14.8–15.5 (15.2±0.3) 13.4–15.7 (14.3±0.6) 0.0513
SL 6.2–6.6 (6.4±0.2) 5.3–6.8 (6.0±0.4) 0.2522
IND 4.2–4.9 (4.5±0.3) 3.3–4.3 (3.9±0.2) 0.0001***
IOD 4.3–4.8 (4.6±0.2) 3.1–3.7 (3.4±0.2) 0.0000***
ED 4.6–5.7 (5.2±0.4) 3.5–5.3 (4.8±0.5) 0.1494
HAL 10.9–12.9 (11.9±0.7) 10.5–13.1 (11.4±0.7) 0.4359
FOL 9.9–11.4 (10.6±0.5) 9.4–11.2 (10.2±0.4) 0.0534
TFL 29.4–31.6 (30.8±0.8) 26.9–32.9 (29.1±1.6) 0.0988
TB 17.6–19.4 (18.7±0.7) 15.9–19.1 (17.4±0.9) 0.0077**
A
A
C
C
E
E
F
F
B
B
D
D
Figure 3 Morphological features of adult male holotype CIB 121683 of Scutiger wolong sp. nov. in life. A: Dorsolateral view; B: Ventral view;
C: Right hand; D: Right foot; E: Close-up of right forelimb, showing black nuptial spines on fingers, large translucent spines on lower arm and
inner upper arm, and small black spines on dorsolateral upper arm; F: Close-up of ventral surface, showing pectoral glands and axillary glands
covered by black dense spines, rough belly rough with black horny spines, and black dense spines on inner forelimb. Photos by Zhitong LYU.
No. 1
Zhitong LYU et al.
A New Species of Scutiger
29

pectoral glands and pair of axillary glands present on chest,
covered by dense spines; axillary gland distinctly isolated from
pectoral gland, about two-thirds of pectoral gland in size;
dense spines on inner forelimb aligned, extending from wrist
to near axilla; belly with scattered horny spines; ventral
hindlimb smooth with several horny spines scattered around
cloaca.
Coloration of holotype In life, dorsal surface brown;
irregular dark brown patches on dorsum and top of head;
dark brown line extending from tip of snout, lower margin of
canthus rostralis, and lower margin of supratympanic fold to
corner of mouth; irregular dark brown patches on lip; various
dark brown stripes on flank and dorsolateral forelimb and
hindlimb; ventral surface with dense net-like stripes; all spines
on the skin black, except for larger and translucent spines on
lower arm and inner upper arm.
Variations and sexual dimorphism Morphometric
differences among specimens are presented in Table 4. All
specimens exhibited similar morphology, except for
differences based on sexual dimorphism: (1) females larger
than males (SVL 54.3–56.3 mm (n=2) in adult females vs. SVL
42.2–46.9 mm (n=6) in adult males) with relatively slender
body; (2) relatively shorter hindlimbs in females (TFL/SVL
0.61–0.62 (n=2) and TB/SVL 0.34–0.36 (n=2) in adult females
vs. TFL/SVL 0.66–0.72 (n=6) and TB/SVL 0.40–0.43 (n=6) in
adult males), with tibio-tarsal articulation reaching shoulder
when hindlimb stretched alongside body; (3) skin smooth
without black horny spines in females; (4) pair of pectoral
glands and pair of axillary glands present on chest in males; (5)
pectoral glands and axillary glands covered by dense spines,
Table 4 Measurements (in mm) of examined specimens of Scutiger chintingensis and S.wolong sp. nov. * for holotype. CIB 121707 was not measured
due to the poor condition of preservation.
Sex Locality SVL HL HW SL IND IOD ED HAL FOL TFL TB
Scutiger chintingensis
CIB 24670* Male Mt. Emei 41.8 14.8 14.3 5.3 4.3 3.4 4.6 11.0 10.1 28.3 17.0
CIB 121687 Male Mt. Emei 37.5 13.8 13.4 5.8 3.6 3.4 3.5 10.8 10.6 28.2 16.3
CIB 121689 Male Mt. Emei 37.6 13.7 13.4 5.4 3.3 3.1 4.4 10.6 9.6 26.9 15.9
CIB 121690 Male Mt. Emei 43.9 14.3 15.1 6.0 3.8 3.4 4.5 11.3 10.5 28.4 18.2
CIB 121691 Male Mt. Emei 42.2 14.0 13.9 6.0 4.3 3.7 5.1 10.6 9.9 27.0 16.6
CIB 121692 Male Mt. Emei 41.1 14.4 14.1 5.7 3.7 3.1 4.5 10.5 10.4 28.5 16.5
CIB 121693 Male Mt. Emei 44.0 15.8 15.7 5.8 4.1 3.6 5.3 11.7 10.6 30.1 18.0
CIB 121688 Male Mt. Wawu 43.0 14.1 14.0 6.3 3.9 3.3 4.9 10.9 9.9 27.4 16.6
CIB 121698 Male Mt. Wawu 42.8 13.6 13.8 5.9 3.9 3.5 5.1 11.5 10.3 30.4 17.6
CIB 121699 Male Mt. Wawu 43.4 13.7 14.2 6.3 3.9 3.6 4.5 11.5 10.1 29.9 17.8
CIB 121701 Male Mt. Wawu 45.0 14.6 15.0 6.8 4.1 3.7 5.2 12.8 10.3 32.9 19.1
CIB 121702 Male Mt. Wawu 47.2 14.0 14.4 6.2 3.8 3.7 4.3 13.1 9.9 31.8 18.8
CIB 121694 Male Mt. Wawu 44.8 14.0 14.7 6.2 3.8 3.2 5.2 11.4 11.2 29.0 16.9
CIB 121695 Male Mt. Wawu 43.6 13.6 14.3 6.3 3.9 3.3 5.1 11.6 9.6 30.3 17.3
CIB 121696 Male Mt. Wawu 44.9 14.7 14.5 6.1 3.8 3.7 4.9 11.8 9.4 28.4 18.0
CIB 121697 Male Mt. Wawu 45.2 14.0 14.2 6.4 4.0 3.5 4.6 11.0 9.9 28.3 17.3
CIB 121700 Male Mt. Wawu 46.0 14.2 14.6 6.3 3.9 3.3 5.2 11.1 10.3 29.6 17.6
CIB 24671 Female Mt. Wawu 53.1 15.9 16.0 6.2 5.1 4.4 5.0 12.6 11.9 31.0 18.6
CIB 121703 Female Mt. Wawu 55.3 16.7 17.1 5.7 4.6 4.0 5.1 13.4 11.3 31.2 18.9
CIB 121704 Female Mt. Wawu 53.9 16.9 17.1 6.0 4.5 3.9 5.5 12.6 11.7 32.0 18.8
Scutiger wolong sp. nov.
CIB 121679 Male Wolong 46.4 15.5 15.3 6.6 4.7 4.7 5.2 12.9 10.2 30.6 19.0
CIB 121680 Male Wolong 44.2 14.6 14.9 6.2 4.2 4.8 5.3 11.5 10.7 31.2 19.1
CIB 121681 Male Wolong 46.7 16.0 15.5 6.2 4.5 4.7 5.7 12.1 10.9 31.4 18.8
CIB 121682 Male Wolong 42.2 14.2 14.8 6.2 4.3 4.5 4.9 10.9 10.6 29.4 17.6
CIB 121683* Male Wolong 46.9 14.9 15.4 6.6 4.9 4.3 5.2 12.1 9.9 31.6 19.4
CIB 121684 Male Wolong 42.5 15.2 15.1 6.3 4.2 4.6 4.6 11.7 11.4 30.8 18.2
CIB 121685 Female Wolong 54.3 16.1 16.3 7.2 5.1 4.5 5.0 14.2 12.4 33.8 19.7
CIB 121686 Female Wolong 56.3 17.9 17.6 7.3 5.2 4.9 5.6 13.8 12.6 34.5 19.4
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belly rough with horny spines, nuptial spines present on dorsal
and lateral surface of first and second fingers and inner side of
third finger, and dense spines on inner forelimb aligned,
extending from wrist to near axilla in breeding males.
Coloration in life and dorsal tubercules also varied among
individuals (Figure 4).
Distribution and habitat Scutiger wolong sp. nov. is
currently known only from a single locality in Wolong
Town of Wenchuan County, at the western edge of the
Sichuan Basin. Surveys conducted by Fan et al. (2024) in
Qiaoqi Town, Baoxing County, near Wolong, failed to observe
any Scutiger individuals resembling this species. This toad
species inhabits a flowing stream at 2 414–2 488 m a.s.l.,
located near the main road (G350) and surrounded by
shrubland. The type locality faces threats from road
construction and tourism. All type specimens were observed
in late May in a small lotic pool near a waterfall within the
stream. All captured males exhibited nuptial spines on their
fingers, although some individuals had shed the dense spines
on their chest, suggesting that the breeding season occurs in
May. According to Fan et al. (2024), only one individual was
observed in June, with none detected in July.
4.2. Referred specimens Eighteen adult males and three
adult females of S.chintingensis.
Holotype CIB 24670 (field number No. 70) (Figure 5), adult
male, collected by C.C. LIU (given as “C.H. LIU” in Liu and Hu
(1960) due to a typographical error) in June 1955 from
Chinting (= Jinding), Mt. Omei (= Mt. Emei, Emeishan City)
(3 050 m a.s.l.), Szechwan (= Sichuan) Province, China.
Other materials CIB 24671, adult female, collected by C.C.
LIU in June 1955 from Bingxue Temple, Jinding; CIB
121703–121704, two adult females, collected by Wenfeng HE
and Zhiyi CHEN on 24 August 2018 from Laoxionggou,
Jinding (29.5238° N, 103.3388° E, 2 914 m a.s.l.); CIB 121687,
121689–121693, 121707 (Figure 6), seven adult males,
A
A
C
C
E
E
F
F
B
B
D
D
Figure 4 Variations in Scutiger wolong sp. nov. in life. A: Dorsolateral view of female paratype CIB 121686; B: Ventral view of female paratype
CIB 121686; C: Dorsolateral view of female paratype CIB 121685; D: Dorsolateral view of male paratype CIB 121684; E: Dorsolateral view of
male paratype CIB 121681; F: Dorsal view of male paratype CIB 121682. Photos by Zhitong LYU.
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Zhitong LYU et al.
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collected by Dihao WU and Zhongliang PENG on 8 June
2019 from Laoxionggou, Jinding (29.5195°–29.5199° N,
103.3275°–103.3289° E; 2 870–2 885 m a.s.l.); CIB 121688,
121698–121699, 121701–121702, five adult males, collected
by Junjie HUANG on 15 June 2020 from Yuanxi (29.6489° N,
102.9517° E, 2 644 m a.s.l), Mt. Wawu, Hongya County,
A
A
C
C
D
D
B
B
10 mm
Figure 5 Morphological features of adult male holotype CIB 24670 of Scutiger chintingensis in preservative. A: Dorsal view; B: Ventral view; C:
Close-up of ventral upper body, showing pectoral glands and axillary glands covered by black dense spines, smooth belly, and black dense spines
on inner forelimb; D: Right foot. Photos by Zhitong LYU and Ke JIANG.
A
A
B
B
C
C
D
D
Figure 6 Variations in Scutiger chintingensis in life. A: Dorsolateral view of male CIB 121707; B: Ventral view of male CIB 121707; C:
Dorsolateral view of male CIB 121687 and uncaptured female; D: Dorsolateral view of uncaptured female. Photos by Dihao WU.
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Sichuan; CIB 121694–121697, 121700, five adult males,
collected by Dihao WU on 24 June 2020 from Yangxi
(29.6429° N, 102.9509° E, 2 498 m a.s.l.), Mt. Wawu.
5. Discussion
This study established that the Scutiger population from
Wolong, Wenchuan County, previously reported as S.
chintingensis (Yu et al., 1983; Fan et al., 2024), represents a
distinct species, morphologically and phylogenetically
distinguishable from all known congeners. With the
description of this new species, the distribution of S.
chintingensis is redefined as more restricted, limited to only
three adjacent localities. Scutiger chintingensis is currently
classified as Endangered by the China Red List of
Biodiversity (Jiang et al., 2021) and is listed as a protected
species by the National Forestry and Grassland
Administration of China (2021). As the Wolong population
was previously included under these evaluations, we
recommend that Scutiger wolong sp. nov. be similarly
classified as Endangered and afforded at least second-class
protection under the wildlife conservation regulations of
China. Surveys are also urgently required in neighboring areas
to investigate and monitor the population status and
fluctuation of this new rare species.
This research underscores the critical role of taxonomy in
biodiversity science. Accurate taxonomic classification is
indispensable for understanding species richness,
distribution, and ecological roles, forming the foundation for
biodiversity assessments and conservation planning. By
distinguishing Scutiger wolong sp. nov. from S.chintingensis,
this study enables more precise biological research and
conservation efforts for both species. The importance of
taxonomic revisions has been further demonstrated in recent
studies on rare and protected amphibians, such as Tylototriton
sini Lyu, Wang, Zeng, Zhou, Qi, Wan & Wang, 2021 vs. T.
asperrimus Unterstein, 1930, Andrias jiangxiensis Lu, Wang,
Chai, Yi, Peng, Murphy, Zhang & Che, 2022 vs. A.davidianus
(Blanchard, 1871), and Hypselotriton oolong Wang, Zeng, Wei &
Lyu, 2024 vs. H.orphicus (Risch, 1983) (Lyu et al., 2021; Chai et
al., 2022; Wang et al., 2024). These studies highlight the
necessity of accurate taxonomy to ensure that rare and
vulnerable species receive effective protection.
The family Megophryidae Bonaparte, 1850, represents a
diverse and ecologically significant group of amphibians
within the Pan-Himalaya fauna. This family exhibits
remarkable species diversity (Hou et al., 2020; Wang et al.,
2022; Lyu et al., 2023b; Sánchez-Vialas et al., 2024), and the
genus Scutiger is notable for its adaptation to high-altitude
environments, with many species evolving within the
Qinghai-Xizang Plateau (Hoffman et al., 2017, 2024). Since
2019, nine new congeners have been described within the
genus Scutiger, representing approximately 30% for the total
recognized species (Frost, 2024; this study). This rapid rate of
discovery suggests a historical underestimation for the species
diversity within this genus. For instance, samples of S.
liupanensis were revealed as paraphyletic in our phylogenetic
analysis (Figure 1), suggesting potential cryptic species.
Nonetheless, our phylogenetic analysis also recovered small
genetic divergence between S.feiliangi and S.ningshanensis,
raising the caution against taxonomic inflation as suggested in
other groups of Megophryidae (Wang et al., 2022; Lyu et al.,
2023b). Future research should prioritize exploring the
diversity, distribution, and speciation mechanism of Scutiger
species. Such studies will enhance our understanding of how
poikilothermal amphibians adapt to extreme alpine habitats,
contributing to broader insights into the evolutionary
processes shaping biodiversity in harsh environments.
Nomenclatural acts registration The electronic
version of this article in portable document format represents
a published work according to the International Commission
on Zoological Nomenclature (ICZN), and hence the new
names contained in the electronic version are effectively
published under that Code from the electronic edition alone
(see Articles 8.5–8.6 of the Code). This published work and
the nomenclatural acts it contains have been registered in
ZooBank, the online registration system for the ICZN. The
ZooBank LSIDs (Life Science Identifiers) can be resolved and
the associated information can be viewed through any
standard web browser by appending the LSID to the prefix
http://zoobank.org/.
Publication LSID: urn:lsid:zoobank.org:pub:3D8A4003-
DCA0-4B7B-9849-DE2094DB0C4E
Scutiger wolong LSID: urn:lsid:zoobank.org:act:DE1D29D2-
625B-47D7-9808-698F7AF6A87E
Acknowledgements We thank Xiaoyi WANG, Lan
ZHAO, Jinlong REN, Zeng WANG, Zhongliang PENG,
Ruwan JIA, Tao XUE, and Yizhou LIU for their help in the
field and laboratory works. We thank the editor and
reviewers for their constructive suggestions on the
manuscript. This work was supported by the National Key
Program of Research and Development, Ministry of Science
and Technology of China (2023YFF1304800), National
Natural Science Foundation of China (32400361, 32200363,
32370498), Sichuan Science and Technology Program
(2024NSFSC1180, 2023NSFSC1155), the Second Tibetan
Plateau Scientific Expedition and Research Program (STEP)
(2019QZKK0501), Taxonomist Position, Chinese Academy of
No. 1
Zhitong LYU et al.
A New Species of Scutiger
33

Sciences (CAS-TAX-24-052, CAS-TAX-24-051), and
Biological Resources Programme, Chinese Academy of
Sciences (KFJ-BRP-017-086, KFJ-BRP-017-65).
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Scientific Editor: Bin WANG
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�How to cite this article:
Lyu Z. T., Gao Z. Y., Huang J. J., Wu D. H., Jiang D. C., Jiang K., Li J. T. 2025. A new species of Scutiger from Sichuan, China,
previously misreported as S.chintingensis (Anura: Megophryidae). Asian Herpetol Res, 16(1): 20–35. DOI: 10.3724/
ahr.2095-0357.2024.0049. CSTR: 32242.14.ahr.2095-0357.2024.0049
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