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Original Article
A new species of the Amolops monticola group (Anura,Ranidae)from
southern Yunnan, China
Shuo Liu
a
, Mian Hou
b
, Mingzhong Mo
c
,YiLu
d
, Jimin Guo
d
, Wen Wang
d
, Wenxiang Zhang
d
,
Dingqi Rao
e
,
y
, Song Li
a
,
f
,
y
,
*
a
Kunming Natural History Museum of Zoology, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan 650223, China
b
College of Continuing (Online) Education, Sichuan Normal University, Chengdu, Sichuan 610066, China
c
Honghe Prefecture Forestry and Grassland Bureau of Yunnan Province, Mengzi, Yunnan 661199, China
d
Yunnan Huanglianshan National Nature Reserve Management and Protection Bureau, Lvchun, Yunnan 662599, China
e
Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan 650201, China
f
Yunnan Key Laboratory of Biodiversity Information, Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming, Yunnan 650201, China
article info
Article history:
Received 31 May 2024
Received in revised form
8 August 2024
Accepted 15 August 2024
Available online xxx
Keywords:
16S
Cascade frog
Morphology
ND2
Taxonomy
abstract
A new species of the Amolops monticola group is described from Yunnan Huanglianshan National Nature
Reserve, southern Yunnan, China. Morphologically, the new species is characterized by a moderate body
size (snoutevent length 38.3e40.8 mm in adult males and 62.3e63.1 mm in adult females), head being
longer than wide, nostril to snout tip distance being slightly smaller than eye to nostril distance, the
presence of pineal body, distinct tympanum, vomerine teeth being invisible in males and weak but
visible in females, distinct dorsolateral folds, the absence of outer metatarsal tubercle, the presence of
vocal sac and nuptial pad in adult males, dorsal surface of head and body being green with no or a few
black dots, and the presence of a few large flat white tubercles on the posterior inferior flank. Genetically,
the uncorrected genetic distance between the new species and other species of the A. monticola group
ranged from 3.0% to 7.6% in 16S ribosomal RNA and from 7.2% to 17.1% in NADH dehydrogenase subunit 2
gene sequences. Currently, the new species is known only from its type locality in Lvchun County,
Honghe Prefecture, Yunnan Province, China.
Ó2024 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing
Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
Introduction
Amolops Cope, 1865 is a type of frogs that is widely distributed
from southern and eastern Himalayas to the Peninsular Malaysia
(Dever et al. 2012;Pham et al. 2019;Wu et al. 2020;Mahony et al.
2022;Tang et al. 2023;Li et al. 2024;Frost 2024). Species of Amo-
lops usually inhabits torrents or waterfalls, and they have an
abdominal sucker in larvae and enlarged digital discs in adults (Fei
et al. 2009;Fei et al. 2012). Amolops currently contains 85 species,
and the species number is continuously increasing (Frost 2024).
This genus was partitioned into ten species groups, namely the
A. chayuensis group, A. daiyunensis group, A. hainanensis group,
A. larutensis group, A. mantzorum group, A. marmoratus group,
A. monticola group, A. ricketti group, A. spinapectoralis group, and A.
viridimaculatus group (Jiang et al. 2021).
Among the ten species groups, the Amolops monticola group is the
most diverse group that comprises 24 species, namely A. adicola
Patel, Garg, Das, Stuart & Biju, 2021,A. akhaorum Stuart, Bain,
Phimmachak & Spence, 2010,A. aniqiaoensis Dong, Rao & Lü, 2005,
A. archotaphus (Inger & Chan-ard, 1997), A. bellulus Liu, Yang, Ferraris
& Matsui, 2000,A. binchachaensis Rao, Hui, Ma & Zhu, 2022 “2020”,
A. chakrataensis Ray, 1992, A. chaochin Jiang, Ren, Lyu & Li, 2021,
A. chunganensis (Pope, 1929), A. compotrix (Bain, Stuart & Orlov,
2006), A. cucae (Bain, Stuart & Orlov, 2006), A. daorum (Bain, Lathrop,
Murphy, Orlov & Ho, 2003), A. deng Jiang, Wang & Che, 2020,
A. kohimaensis Biju, Mahony & Kamei, 2010, A. iriodes (Bain &
Nguyen, 2004), A. mengdingensis Yu, Wu & Yang, 2019,
A. mengyangensis Wu & Tian, 1995, A. monticola (Anderson, 1871),
A. nyingchiensis Jiang, Wang, Xie, Jiang & Che, 2016,A. putaoensis Gan,
Qin, Lwin, Li, Quan, Liu & Yu, 2020, A. truongi Pham, Pham, Ngo, Sung,
Ziegler & Le, 2023, A. tuanjieensis Gan, Yu & Wu, 2020, A. vitreus (Bain,
*Corresponding author. ORCID.: 0000-0003-0834-5882
E-mail addresses: raodq@mail.kiz.ac.cn (D. Rao), lis@mail.kiz.ac.cn (S. Li).
Peer review under responsibility of National Science Museum of Korea (NSMK) and
Korea National Arboretum (KNA).
y
The authors contributed equally to this article.
Contents lists available at ScienceDirect
Journal of Asia-Pacific Biodiversity
journal homepage: http://www.elsevier.com/locate/japb
https://doi.org/10.1016/j.japb.2024.08.002
pISSN2287-884X eISSN2287-9544/Ó2024 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open
access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
Stuart & Orlov, 2006), and A. wenshanensis Yuan, Jin, Li, Stuart & Wu,
2018 (Jiang et al. 2021;Pham et al. 2023; Frost 2024;Li et al. 2024).
During our recent field work in southern Yunnan, China, from
April to July 2023, some specimens of Amolops were collected from
Yunnan Huanglianshan National Nature Reserve. Molecular and
morphological comparison revealed that these specimens belong
to a distinct taxon in the A. monticola group. Herein, we describe
them as a new species of Amolops.
Material and methods
Field surveys were conducted in Yunnan Huanglianshan Na-
tional Nature Reserve, Lvchun County, Honghe Prefecture, Yunnan
Province, China, under the permit from Yunnan Huanglianshan
National Nature Reserve Management and Protection Bureau.
Specimens were preserved in 75% ethanol. Liver tissues were stored
in analytical pure ethanol. The newly collected specimens were
deposited at Kunming Natural History Museum of Zoology,
Kunming Institute of Zoology, Chinese Academy of Sciences (KIZ).
Total genomic DNA was extracted from tissues. Fragments of
mitochondrial 16S ribosomal RNA (16S) and NADH dehydrogenase
subunit 2 (ND2) genes were amplified and sequenced. The primer
pair L2188: 5
0
eAAAGTGGGCCTAAAAGCAGCCAe3
0
and 16H1: 5
0
e
CTCCGGTCTGAACTCAGATCACGTAGGe3
0
(Hedges 1994;Matsui
et al. 2006) was used for 16S, and the primer pair Met-LND2: 5
0
e
e3
0
(Stuart et al. 2006) was used for ND2. The amplification and
sequencing were completed by Tsingke Biotechnology Co., Ltd. The
newly generated sequences have been deposited in GenBank, and
homologous sequences were obtained from GenBank (Table 1).
Sequences were aligned using MAFFT 7.471 (Katoh and Standley
2013). Phylogenetic analyses were constructed based on the
concatenated sequences of 16S and ND2 genes. The best fit sub-
stitution models were selected under the Akaike Information
Criterion in ModelFinder (Kalyaanamoorthy et al. 2017). Bayesian
inference (BI) was performed in MrBayes 3.2.7 (Ronquist et al.
2012) using the GTR þFþIþG4 model, and the Markov chains
were run for 1,000,000 generations and sampled every 100 gen-
erations. Maximum likelihood (ML) analysis was performed in IQ-
TREE 1.6.12 (Nguyen et al. 2015) using the TPM2u þFþR3
model, and the branch support was assessed based on 1,000 ul-
trafast bootstrap replicates. The uncorrected pairwise distances
between species were calculated for 16S and ND2 separately in
MEGA 11 (Tamura et al. 2021).
Measurements were taken with a digital caliper to the nearest
0.1 mm. Measurement methodology followed Pham et al. (2019).
The following morphological characteristics were noted: eye
diameter (EYE); finger disk width (FPW), at the widest part of the
pad of finger III; head length (HDL), from the tip of the snout to the
articulation of the jaw; head width (HDW), between left and right
articulations of the quadratojugal and maxilla; interorbital distance
(IOD); snout length (SNT); snoutevent length (SVL); tympanume
Table 1. Samples used for the phylogenetic analyses in this study.
Species Voucher Locality 16S ND2
Amolops huanglianshanensis sp. nov. KIZ2023089 China: Lvchun, Yunnan PQ202844 PQ210059
Amolops huanglianshanensis sp. nov. KIZ2023090 China: Lvchun, Yunnan PQ202845 PQ210060
Amolops huanglianshanensis sp. nov. KIZ2023091 China: Lvchun, Yunnan PQ202846 PQ210061
Amolops huanglianshanensis sp. nov. KIZ2023092 China: Lvchun, Yunnan PQ202847 PQ210062
Amolops huanglianshanensis sp. nov. KIZ2023093 China: Lvchun, Yunnan PQ202848 PQ210063
Amolops huanglianshanensis sp. nov. KIZ2023094 China: Lvchun, Yunnan PQ202849 PQ210064
Amolops huanglianshanensis sp. nov. KIZ2023095 China: Lvchun, Yunnan PQ202850 PQ210065
Amolops huanglianshanensis sp. nov. KIZ2023097 China: Lvchun, Yunnan PQ202851 PQ210066
Amolops huanglianshanensis sp. nov. KIZ2023098 China: Lvchun, Yunnan PQ202852 PQ210067
Amolops huanglianshanensis sp. nov. KIZ2023099 China: Lvchun, Yunnan PQ202853 PQ210068
Amolops adicola BNHS 6121 India: Arunachal Pradesh MZ229772 MZ231116
Amolops akhaorum FMNH 271355 Laos: Luang Namtha FJ417158 FJ417207
Amolops aniqiaoensis SYNU 04II6015 China: Medog, Tibet MN953655 MN958714
Amolops archotaphus FMNH 271708 Thailand: Chiang Mai MN953659 MN958718
Amolops cf. bellulus CAS 233991 China: Tengchong, Yunnan FJ417127 FJ417176
Amolops chaochin SCUM045818HX China: Anxian, Sichuan MN953669 FJ417179
Amolops cucae AMNH 168727 Vietnam: Van Ban, Lao Cai FJ417144 FJ417193
Amolops compotrix FMNH 256500 Laos: Nakai, Khammouan FJ417141 FJ417190
Amolops chunganensis KIZYPX18652 China: Chengkou, Chongqing MN953670 MN958728
Amolops daorum ROM 38501 Vietnam: Sa Pa, Lao Cai FJ417150 FJ417199
Amolops daorum ROM 38503 Vietnam: Sa Pa, Lao Cai FJ417151 FJ417200
Amolops daorum MNHN 1999.5811 Vietnam: Sa Pa, Lao Cai KR827703 /
Amolops daorum MNHN 1999.5812 Vietnam: Sa Pa, Lao Cai KR827704 /
Amolops cf. daorum FMNH 255353 Laos: Vieng Tong, Huaphahn FJ417147 FJ417196
Amolops cf. daorum FMNH 255354 Laos: Vieng Tong, Huaphahn FJ417148 FJ417197
Amolops cf. daorum FMNH 255355 Laos: Vieng Tong, Huaphahn FJ417149 FJ417198
Amolops deng KIZ 014116 China: Zayü, Tibet MN953695 MN958752
Amolops iriodes AMNH 163925 Vietnam: Vi Xuyen, Ha Giang FJ417154 FJ417203
Amolops iriodes AMNH 163926 Vietnam: Vi Xuyen, Ha Giang FJ417152 FJ417201
Amolops iriodes AMNH 163928 paratype Vietnam: Vi Xuyen, Ha Giang FJ417153 FJ417202
Amolops kohimaensis WIIADA 751 India: Nagaland MZ229774 MZ231118
Amolops mengdingensis KIZ 20160266 China: Mengding, Yunnan MK501809 MK501815
Amolops monticola WIIADA 544 India: South Sikkim MZ229773 MZ231117
Amolops nyingchiensis KIZ 016415 China: Medog, Tibet MN953718 MN958776
Amolops putaoensis GXNU W011 Myanmar: Putao, Kachin MT901383 MT901213
Amolops truongi ZVNU.2022.01 Vietnam: Muong La, Son La / OP157200
Amolops tuanjieensis GXNU YU110003 China: Tuanjie, Yunnan MN832772 MN832756
Amolops tuberodepressus SCUM050433CHX China: Jingdong, Yunnan MN953729 MN958786
Amolops viridimaculatus KIZ 048487 China: Tengchong, Yunnan MN953731 MN958788
Amolops vitreus FMNH 258183 Laos: Phongsaly, Phongsaly FJ417163 FJ417212
Amolops wenshanensis KIZ 021425 China: Xichou, Yunnan MN953724 MG996763
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx2
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
eye distance (TEY); tibial length (TIB); tympanum diameter (TMP);
and toe disk width (TPW), at the widest part of the pad of toe III.
We compared morphological characters of the new species with
all other members of the Amolops monticola group relying on
original species descriptions (Wu and Tian 1995;Inger and Chan-
ard 1997;Liu et al. 2000;Bain et al. 2003,2006;Bain and Nguyen
2004;Zhao et al. 2005;Biju et al. 2010;Stuart et al. 2010;Jiang
et al. 2016,2021;Yuan et al. 2018;Yu et al. 2019;Che et al. 2020;
Gan et al. 2020a,2020b;Patel et al. 2021;Rao 2022“2020”;Pham
et al. 2023) and the additional data from Bain et al. (2003,2006),
Stuart et al. (2010),Jiang et al. (2021), and Patel et al. (2021).
Results
The BI and ML analyses yielded consistent topology. All se-
quences of the newly collected specimens clustered together and
represented a distinct lineage in the Amolops monticola group. The
novel lineage was recovered as the sister clade to the clade con-
sisting of A. iriodes,A. daorum, and A. cf. daorum, but the support
rates were very low by both BI and ML (Figure 1). The uncorrected
pairwise distances between the sequences of the newly collected
specimens and the sequences of other species in the A. monticola
group ranged from 3.0% to 7.6% in 16S (Table 2) and from 7.2% to
17.1% in ND2 (Table 3).
Amolops huanglianshanensis sp. nov.
(Figures 2e5)
LSID: urn:lsid:zoobank.org:act:D5DA4C33-0EA9-4890-B4D1-
38FE2BD42962
Type. Holotype. KIZ2023094, adult male, collected on 28 April
2023 by Shuo Liu from Sanmeng Township, Lvchun County, Honghe
Prefecture, Yunnan Province, China (22
53
0
21
00
N, 102
17
0
44
00
E;
1800 m).
Figure 1. Bayesian phylogram of the Amolops monticola group inferred from the combination of 16S rRNA and ND2 sequences. Numbers after and behind “/”are Bayesian posterior
probabilities and ML bootstrap values (only values above 0.90/90 are shown), respectively.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 3
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
Paratypes. KIZ2023089eKIZ2023093, five adult males, and
KIZ2023095, adult female, the same collection data as the holotype.
KIZ2023097, adult male, and KIZ2023098eKIZ2023099, two adult
females, collected on 18 July 2023 by Shuo Liu from the same lo-
cality as the holotype.
Diagnosis. The new species is assigned to genus Amolops based
on the presence of enlarged digital discs, circummarginal groove on
the discs of fingers, well-developed webbing between toes, and the
absence of webbing between fingers. It is further assigned to the
A. monticola group based on having smooth skin, side of the head
dark with a light-colored upper lip stripe extending to the axilla,
and distinct dorsolateral folds. It can be distinguished from other
species in the A. monticola group by having a combination of the
following characters: Body size moderate, SVL 38.3e40.8 mm in
adult males and 62.3e63.1 mm in adult females; head longer than
wide; nostril to snout tip distance slightly smaller than eye to
nostril distance; internarial distance greater than interorbital dis-
tance in males and internarial distance approximately equal to
interorbital distance in females; upper eyelid width approximately
equal to interorbital distance in males and upper eyelid width
smaller than interorbital distance in females; pineal body present;
vomerine teeth invisible in males and weak but visible in females;
tympanum distinct, small; supratympanic fold indistinct; inner
metatarsal tubercle small; outer metatarsal tubercle absent; vocal
sac present in adult males; nuptial pad present on finger I of adult
males. Dorsal surface of head and body green with no or a few black
dots; dorsolateral folds pinkish brown; a few large flat white tu-
bercles on posterior inferior flank.
Description of holotype.Adult male, SVL 40.8 mm; head longer than
wide (HDL/HDW 1.12); snoutmoderate long (SNT/HDL0.46), obtusely
Table 2. Uncorrected pairwise genetic distance (%) between members of the Amolops monticola group estimated from 16S sequences.
123456789101112131415161718192021
1Amolops huanglianshanensis sp. nov.
2Amolops adicola 5.6
3Amolops akhaorum 5.3 7.1
4Amolops aniqiaoensis 4.0 4.9 5.8
5Amolops archotaphus 3.0 7.9 4.3 5.8
6Amolops cf. bellulus 5.7 5.3 8.6 4.9 5.6
7Amolops chaochin 3.8 6.6 6.2 5.1 4.9 4.3
8Amolops chunganensis 5.3 9.1 6.4 6.6 6.4 6.0 4.9
9Amolops compotrix 6.2 6.4 7.3 6.0 4.9 7.2 4.5 5.7
10 Amolops cucae 6.8 5.8 7.7 5.8 5.6 7.3 4.5 5.5 2.4
11 Amolops daorum 3.2 6.8 5.7 5.5 4.0 6.3 4.3 5.3 5.9 5.8
12 Amolops cf. daorum 3.6 7.0 5.5 4.8 3.5 6.6 3.6 3.8 5.8 6.1 2.2
13 Amolops deng 3.2 5.3 5.7 4.2 5.1 2.1 4.2 5.5 4.4 4.3 4.6 3.5
14 Amolops iriodes 3.5 6.4 5.9 4.7 3.4 6.7 3.3 4.5 6.3 6.3 0.7 2.2 3.6
15 Amolops kohimaensis 3.4 4.5 5.6 3.4 5.6 2.6 3.6 5.7 4.2 4.0 4.8 4.3 2.3 4.0
16 Amolops mengdingensis 4.5 6.2 4.3 4.5 2.4 7.5 4.3 5.5 6.7 7.5 4.6 4.4 3.8 4.9 4.3
17 Amolops monticola 6.4 5.5 7.9 7.0 7.8 7.4 7.2 9.0 8.0 7.8 7.0 7.1 7.0 6.2 5.8 7.0
18 Amolops nyingchiensis 3.0 5.6 5.8 4.5 5.3 1.1 4.0 6.0 4.5 4.5 4.5 3.8 1.7 3.6 2.3 4.0 7.0
19 Amolops putaoensis 3.6 5.2 6.1 4.2 5.0 2.5 4.2 6.2 4.2 4.0 4.2 3.6 2.4 3.5 2.2 4.6 6.6 2.0
20 Amolops tuanjieensis 3.4 6.4 5.6 4.2 3.4 6.1 3.8 4.9 5.9 6.7 3.7 3.3 2.3 3.7 3.4 4.2 7.2 2.6 3.2
21 Amolops vitreus 7.6 4.0 8.5 1.5 4.0 8.8 1.0 2.5 3.0 3.6 5.6 6.2 2.5 7.3 2.0 8.3 5.2 3.0 1.8 6.6
22 Amolops wenshanensis 4.2 6.8 5.7 6.0 4.3 4.5 4.5 6.1 2.3 2.6 4.6 3.7 4.4 3.6 4.9 3.4 7.9 4.5 4.0 4.2 2.0
Table 3. Uncorrected pairwise genetic distance (%) between members of the Amolops monticola group estimated from ND2 sequences.
1 2 3 4 5 6 7 8 9 10111213141516171819202122
1Amolops huanglianshanensis
sp. nov.
2Amolops adicola 14.8
3Amolops akhaorum 11.5 17.4
4Amolops aniqiaoensis 14.7 8.1 14.8
5Amolops archotaphus 11.7 17.6 13.1 17.1
6Amolops cf. bellulus 14.8 10.9 16.6 11.7 16.3
7Amolops chaochin 15.5 13.1 17.7 14.6 18.7 16.0
8Amolops chunganensis 17.1 12.6 16.4 14.8 19.8 15.5 13.1
9Amolops compotrix 16.0 15.0 17.4 15.3 18.2 15.7 15.8 17.9
10 Amolops cucae 16.5 15.7 16.9 15.4 16.5 15.7 17.5 17.5 7.7
11 Amolops daorum 7.2 16.9 12.1 15.9 12.2 15.9 16.7 16.9 17.1 17.7
12 Amolops cf. daorum 7.5 15.2 11.6 15.1 12.1 15.4 16.6 15.8 17.3 18.5 4.0
13 Amolops deng 13.3 12.9 15.5 12.8 16.3 8.2 16.2 15.0 16.3 14.5 14.2 13.8
14 Amolops iriodes 8.0 17.1 12.4 15.7 13.0 15.9 16.8 16.3 17.0 17.4 2.8 4.8 14.0
15 Amolops kohimaensis 14.5 7.0 15.6 6.5 16.4 9.4 15.1 14.7 14.8 14.7 14.7 14.0 10.4 14.9
16 Amolops mengdingensis 12.5 18.5 14.1 16.4 12.9 17.7 19.3 17.6 18.4 18.2 13.9 13.7 16.8 14.1 17.1
17 Amolops monticola 16.6 8.3 16.3 8.2 17.5 11.2 15.2 16.1 15.8 15.7 16.1 15.8 12.3 16.0 8.2 17.5
18 Amolops nyingchiensis 14.8 12.6 16.4 11.7 16.6 7.9 15.7 15.8 15.8 16.0 14.9 14.8 8.2 14.5 10.1 17.3 11.0
19 Amolops putaoensis 15.8 11.5 15.9 10.2 17.0 11.0 16.5 16.6 16.2 15.7 15.8 15.2 12.6 15.3 9.7 18.0 9.3 11.6
20 Amolops truongi 15.9 15.2 16.6 14.7 18.2 15.6 16.3 17.8 3.5 7.5 16.5 16.9 15.7 16.2 14.6 17.9 15.8 15.6 16.3
21 Amolops tuanjieensis 10.7 17.2 13.1 15.2 12.7 15.2 17.1 16.7 16.5 16.5 9.6 9.8 15.3 10.3 13.9 15.0 14.5 14.3 14.8 16.7
22 Amolops vitreus 14.8 14.4 15.6 14.6 16.2 14.9 15.7 16.6 11.2 12.1 16.1 15.7 14.3 16.1 13.4 18.5 14.0 15.3 14.7 11.5 14.8
23 Amolops wenshanensis 14.9 15.2 14.9 14.3 15.8 15.0 15.6 16.4 7.0 6.9 15.7 15.5 15.4 14.9 14.0 16.9 13.9 14.6 14.7 6.7 14.5 10.2
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx4
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
pointed, rounded in profile, projecting beyond margin of lower jaw in
ventralview; canthus rostralis distinct; loreal region slightly concave;
nostril closer to snout tipthan to eye; pinealbody present; internarial
distance greater than interorbital distance; upper eyelid width
approximately equal to interorbital distance; pupil oval, horizontal;
tympanum distinct, small (TMP/HDL 0.13); tympanum-eye distance
smaller than tympanum diameter; supratympanic fold indistinct;
vomerine teeth invisible; choanae rounded; tongue cordiform,
notched posteriorly; paired vocal sacs at angles of jaw.
Forelimb relatively long; relative length of fingers
III >IV >II >I; tips of outer three fingers significantly expanded
into discs, tip of first finger slightly expanded into disc, all discs
with circummarginal grooves; nuptial pad present on finger I;
webbing between fingers absent; subarticular tubercles present,
oval, formula 1, 1, 2, 2; supernumerary tubercles absent; inner
metacarpal (thenar) tubercle large, oval; outer metacarpal tubercle
indistinct.
Hind limb long; TIB/SVL 0.63; relative length of toes
IV >V>III >II >I; all toe tips expanded into discs with circum-
marginal grooves; webbing between toes deeply incurved,
webbing formula I0e1/2II0e1III0e1IV1e0V; subarticular tubercles
distinct, oval, formula 1, 1, 2, 3, 2; inner metatarsal tubercle small,
oval; outer metatarsal tubercle absent; tarsal fold absent.
Doral skin smooth, dorsolateral folds distinct, flank skin smooth
with a few large flat white tubercles on posterior inferior region;
ventral skin smooth with many small flat tubercles on basal ventral
thigh.
Color of holotype in life. Dorsal surface of head and body green
with several black dots; dorsolateral folds pinkish brown; lateral
surface head brownish black; a white stripe extending along upper
lip to shoulder on each side; upper part of iris light brown, lower
Figure 2. Dorsal view (top) and ventral view (bottom) of the type series of Amolops huanglianshanensis sp. nov. in preservative.
Figure 3. Close-up views of the hand and foot of the holotype (KIZ2023094) of
Amolops huanglianshanensis sp. nov. in preservative: A, volar view of the left hand; B,
plantar view of the left foot.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 5
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
part of iris dark brown; tympanum dark brown; upper flank
brownish black, middle flank green, lower flank brown with some
white spots; dorsal surface of fore and hind limbs light brown with
brownish black bands; ventral surface of head and body white;
vocal sac light flesh colored; ventral surface of fore limb light flesh
colored, ventral surface of hind limb light flesh colored with some
grayish brown spots; toe webbing brownish yellow with some dark
brown spots.
Variations. The females have a larger body size than the males;
the internarial distance is approximately equal to the interorbital
distance, the upper eyelid width is smaller than the interorbital
distance, and the tympanum to eye distance is approximately equal
to the tympanum diameter in the females (Table 4); in addition, the
vomerine teeth are weak but visible in the females. Most males
have several black dots on dorsum, a few males and all females
have no black dot on dorsum, and there are some brown blotches
on the throat and chest in a few individuals.
Ecology notes. The specimens of the new species were found on
the leaves of tall herbaceous plants beside a river at night (Figure 6).
No courtship calls were heard and no amplexus behavior and
tadpoles were found.
Distribution. The new species is currently known only from
Yunnan Huanglianshan National Nature Reserve in Lvchun County,
Honghe Prefecture, Yunnan Province, China (Figure 7).
Etymology. The specific epithet is named for the type locality
Yunnan Huanglianshan National Nature Reserve. We suggest the
Figure 4. Dorsal (A), lateral (B), and ventral (C) views of the male holotype (KIZ2023094), and dorsal (D), lateral (E), and ventral (F) views of the female paratype (KIZ2023099) of
Amolops huanglianshanensis sp. nov. in life.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx6
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
Figure 5. Dorsolateral (A) and ventral (B) views of the male paratype (KIZ2023092), dorsolateral (C) and ventral (D) views of the male paratype (KIZ2023093), and dorsolateral (E)
and ventral (F) views of the female paratype (KIZ2023098) of Amolops huanglianshanensis sp. nov. in life.
Table 4. Measurements (mm) of the type series of Amolops huanglianshanensis sp. nov.
KIZ2023094
Male holotype
KIZ2023089
Male paratype
KIZ2023090
Male paratype
KIZ2023091
Male paratype
KIZ2023092
Male paratype
KIZ2023093
Male paratype
KIZ2023097
Male paratype
KIZ2023095
Female paratype
KIZ2023098
Female paratype
KIZ202399
Female paratype
SVL 40.8 39.1 40.7 38.7 38.8 38.3 39.9 63.1 62.3 62.6
HDL 15.0 14.0 14.9 13.5 13.6 14.1 14.0 21.4 21.9 21.2
HDW 13.4 12.8 14.3 12.6 12.6 13.1 12.9 19.8 20.7 19.8
SNT 6.9 6.5 6.8 6.1 6.4 6.4 6.3 9.9 10.1 10.2
EYE 5.5 5.1 5.5 5.0 5.1 5.2 5.3 7.0 7.0 7.1
IOD 3.5 3.3 3.6 3.5 3.5 3.4 3.3 6.2 6.1 6.3
TMP 2.0 2.0 2.0 1.9 1.9 1.9 2.0 2.2 2.4 2.4
TEY 1.3 1.2 1.2 1.2 1.2 1.2 1.3 2.2 2.3 2.4
FPW 2.5 2.2 2.4 2.2 2.3 2.2 2.4 3.5 3.7 3.8
TPW 1.9 1.7 1.9 1.6 1.8 1.7 1.7 2.6 2.9 2.8
TIB 25.7 24.4 24.9 24.7 23.6 23.8 24.7 40.9 39.0 38.6
SVL ¼snoutevent length; HDL ¼head length; HDW ¼head width; SNT ¼snout length; EYE ¼eye diameter; IOD ¼interorbital distance; TMP ¼tympanum diameter;
TEY ¼tympanumeeye distance; FPW ¼finger disk width; TPW ¼toe disk width; TIB ¼tibial length.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 7
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
English common name “Huanglianshan cascade frog”and the
Chinese common name “黄连山湍蛙(Pinyin: huáng lián sh
an tu
an
w
a)”.
Comparisons. Amolops huanglianshanensis sp. nov. closely re-
sembles A. daorum and A. iriodes, however, Amolops huanglian-
shanensis sp. nov. can be distinguished from A. daorum by having a
larger body size (SVL 38.3e40.8 mm vs. 32.0e38.1 mm in males and
62.3e63.1 mm vs. 53.3e57.6 mm in females), a relatively longer
snout (SNT/HDL 0.45e0.47 vs. 0.23e0.24 in males and 0.46e0.48 vs.
0.26e0.30), relatively larger eyes (EYE/HDL 0.36e0.38 vs. 0.20e
0.25 in males and 0.32e0.33 vs. 0.11e0.12), a relatively greater
interorbital distance (IOD/HDL 0.23e0.26 vs. 0.17e0.20 in males
and 0.28e0.30 vs. 0.14e0.21), a smaller ratio of head length to head
width in males (HDL/HDW 1.04e1.12 vs. 1.30e1.66), and a smaller
ratio of tympanum diameter to eye diameter in females (TMP/EYE
0.31e0.34 vs. 0.43e0.47); Amolops huanglianshanensis sp. nov. can
be distinguished from A. iriodes by having a relatively longer snout
in males (SNT/HDL 0.45e0.47 vs. 0.36e0.42), relatively larger eyes
in males (EYE/HDL 0.36e0.38 vs. 0.22e0.27), a relatively smaller
interorbital distance in males (IOD/HDL 0.23e0.26 vs. 0.28e0.32), a
smaller ratio of tympanum diameter to eye diameter in males
(TMP/EYE 0.36e0.39 vs. 0.51e0.63), and a larger ratio of eye
diameter to snout length in males (EYE/SNT 0.78e0.84 vs. 0.57e
0.69) (see Table 5).
Amolops huanglianshanensis sp. nov. can be distinguished from
A.adicola by having a smaller body size in males (SVL 38.3e
40.8 mm vs. 44.9e47.0 mm) and different dorsal color (green vs.
light brown or greyish brown); from A.akhaorum by having a larger
body size in males (SVL 38.3e40.8 mm vs. 34.9e37.2 mm) and no or
a few black dots on dorsum (vs. many black dots on dorsum); from
A.aniqiaoensis by having a smaller body size in males (SVL 38.3e
40.8 mm vs. ca. 52.0 mm) and different dorsal color (green vs. olive
or brown); from A.archotaphus by the absence of outer metatarsal
tubercle (vs. presence) and dorsolateral fold distinct (vs. indistinct
or absent); from A.bellulus by having a smaller body size in males
(SVL 38.3e40.8 mm vs. 45.9e50.1 mm), the presence of vocal sac in
males (vs. absence), and different dorsal color (green vs. sandy
beige); from A. binchachaensis by having a smaller body size (SVL
38.3e40.8 mm vs. ca. 50 mm in males and 62.3e63.1 mm vs. ca.
65 mm in females), head longer than wide (vs. head length
approximately equal to head width), the presence of vocal sac in
males (vs. absence) and different dorsal color (green vs. yellow);
from A.chakrataensis by having a larger body size in females (SVL
62.3e63.1 mm vs. 55.0 mm), head longer than wide (vs. head wider
than long), and different dorsal color (green vs. reddish brown);
from A. chaochin by having a larger body size in females (SVL 62.3e
63.1 mm vs. 50.5e54.4 mm) and different dorsal color (green vs.
brown); from A.chunganensis by having a larger body size in fe-
males (SVL 62.3e63.1 mm vs. 52.6 mm) and different dorsal color
(green vs. pale brown); from A.compotrix by having a larger body
size in females (SVL 62.3e63.1 mm vs. 55.6e56.9 mm) and the
absence of outer metatarsal tubercle (vs. presence); from A.cucae
by having a smaller body size (SVL 38.3e40.8 mm vs. 40.7e44.6 mm
in males and 62.3e63.1 mm vs. 65.8e68.0 mm in females) and the
absence of outer metatarsal tubercle (vs. presence); from A.deng by
having a smaller body size (SVL 38.3e40.8 mm vs. 50.3e57.6 mm in
males and 62.3e63.1 mm vs. 68.5e72.0 mm in females) and
different dorsal color (green vs. brown); from A.kohimaensis by
having a smaller body size in males (SVL 38.3e40.8 mm vs. 42.8e
48.6 mm) and different dorsal color (green vs. brown); from A.
mengdingensis by having a smaller body size in females (SVL 62.3e
63.1 mm vs.. 64.3 mm) and grayish brown spots on ventral hind
limbs (vs. no spots); from A.mengyangensis by tibiotarsal articula-
tion reaching beyond snout tip (vs. reaching snout tip or between
eye and snout tip) and the second finger longer than the first finger
(vs. the first and second fingers are approximately equal in length);
from A. monticola by relative length of toes IV >V>III >II >I(vs.
IV >III >V>II >I) and different dorsal color (green vs. light to dark
brown, reddish brown or ash greyish brown); from A.nyingchiensis
by having a smaller body size in males (SVL 38.3e40.8 mm vs.
48.5e58.3 mm), the presence of vocal sac in males (vs. absence) and
different dorsal color (green vs. brown); from A.putaoensis by head
longer than wide (vs. head wider than long) and different dorsal
color (green vs. brown); from A. truongi by the presence of cir-
cummarginal groove on tip of first finger (vs. absence) and rela-
tively larger disc on tip of the third finger (FPW/SVL 0.06 vs. 0.03e
0.05); from A.tuanjieensis by having a larger body size in females
(SVL 62.3e63.1 mm vs. 56.8e60.7 mm) and different dorsal color
(green vs. reddish brown); from A. vitreus by the absence of outer
metatarsal tubercle (vs.. presence) and visible pineal body (vs. no
visible pineal body); from A.wenshanensis by having a larger size in
females (SVL 62.3e63.1 mm vs. 43.7e45.6 mm) and the presence of
pineal body (absence).
Discussion
Grosjean et al. (2015) identified two specimens (MNHN
1999.5811e1999.5812) from Sa Pa, Lao Cai, Vietnam, as Amolops
mengyangensis and deposited the gene sequences of these two
specimens on GenBank (KR827703eKR827704). In this study, the
sequences of these two specimens were clustered with the se-
quences of the two paratypes (ROM 38501, 38503) of A. daorum,
and the genetic distance between them was only 0.06% in 16S gene.
The type locality of A. daorum is in Sa Pa, Lao Cai, Vietnam, while the
type locality of A. mengyangensis is in Mengyang Town, Jinghong
City, Xishuangbanna Prefecture, Yunnan Province, China (Wu and
Tian 1995;Bain et al. 2003). The two localities are approximately
Figure 6. The habitat at the type locality of Amolops huanglianshanensis sp. nov.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx8
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
300 km apart (Figure 7). Therefore, we consider that the two
specimens (MNHN 1999.5811e1999.5812) should belong to
A. daorum rather than A. mengyangensis.
In the phylogenetic analysis, Amolops daorum and A. iriodes
formed sister groups with strong support, and the genetic distance
between them was very small (0.7% in 16S and 2.8% in ND2). From
this perspective alone, A. daorum and A. iriodes are likely to be
conspecific. However, according to Bain et al. (2003,2006) and
Stuart et al. (2010),A. daorum and A. iriodes have distinct
morphological differences. Therefore, we currently consider that
A. daorum and A. iriodes are not conspecific, but further research is
needed to address the relationship between them. In addition,
Stuart et al. (2010) identified three specimens (FMNH 255353e
255355) from Laos as A. daorum as they have no morphological
difference from A. daorum. However, these three specimens were
not clustered with A. daorum in the phylogenetic analysis. There-
fore, they may represent a cryptic species and we refer to them as
Amolops cf. daorum in this study. Perhaps more research methods
Figure 7. Map showing the type localities of Amolops huanglianshanensis sp. nov. (black star), A. daorum (black dot), A. iriodes (black triangle), and A . mengyangensis (black square).
Table 5. Morphological comparisons among Amolops huanglianshanensis sp. nov., A. daorum, and A. iriodes.
Amolops huanglianshanensis sp. nov. Amolops daorum Amolops iriodes
Males (n¼7) Females (n¼3) Males (n¼7) Females (n¼9) Males (n¼3) Female (n¼1)
SVL 38.3e40.8 62.3e63.1 32.0e38.1 53.3e57.6 38.8e43.2 61.9
HDL/SVL 0.35e0.37 0.34e0.35 / / 0.32e0.39 0.36
HDL/HDW 1.04e1.12 1.06e1.08 1.30e1.66 0.80e1.12 0.99e1.12 1.09
SNT/HDL 0.45e0.47 0.46e0.48 0.23e0.24 0.26e0.30 0.36e0.42 0.37
EYE/HDL 0.36e0.38 0.32e0.33 0.20e0.25 0.11e0.12 0.22e0.27 0.35
TMP/HDL 0.13e0.14 0.11 / / 0.14 0.12
EYE/SNT 0.78e0.84 0.69e0.71 / / 0.57e0.69 0.94
TMP/EYE 0.36e0.39 0.31e0.34 0.19e0.38 0.43e0.47 0.51e0.63 0.35
IOD/HDL 0.23e0.26 0.28e0.30 0.17e0.20 0.14e0.21 0.28e0.32 0.25
TIB/SVL 0.61e0.64 0.62e0.65 0.52e0.68 0.62e0.63 0.59e0.68 0.59
SVL ¼snoutevent length; HDL ¼head length; HDW ¼head width; SNT ¼snout length; EYE ¼eye diameter; IOD ¼interorbital distance; TMP ¼tympanum diameter;
TEY ¼tympanumeeye distance; FPW ¼finger disk width; TPW ¼toe disk width; TIB ¼tibial length.
Data for A. daorum, and A. iriodes were obtained from Bain et al (2003) and Bain and Nguyen (2004), respectively.
S Liu et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 9
Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002
are needed, such as comparing bones, courtship calls, and repro-
ductive habits, to effectively distinguish them from A. daorum.
Yunnan Huanglianshan National Nature Reserve is located in
southern Yunnan Province, between 102
5
0
26
00
e102
25
0
7
00
E and
22
33
0
34
00
e22
59
0
20
00
N, with an altitude span from 320 m to
2637 m (Xu 2003). Previously, two species of Amolops were recor-
ded in this nature reserve, namely A. truongi Pham, Pham, Ngo,
Sung, Ziegler & Le, 2023 and A. viridimaculatus (Jiang, 1983)
(AmphibiaChina 2024;Liu et al. 2024). In addition to the new
species described in this study, we also discovered A. vitreus during
the field surveys in this nature reserves. Therefore, so far, there are
four species of Amolops distributed in this nature reserve, namely
Amolops huanglianshanensis sp. nov., A. truongi,A. viridimaculatus,
and A. vitreus. Within this nature reserve, A. viridimaculatus is
distributed in high-altitude areas, usually above 2000 m; Amolops
huanglianshanensis sp. nov. is distributed in slightly lower areas,
around 1800 m; A. truongi is distributed in moderate-altitude areas,
around 1400 m; and A. vitreus is distributed in low-altitude areas,
below 700 m. No sympatric distribution of these four species was
found. Yunnan Huanglianshan National Nature Reserve has a vari-
ety of terrain and vegetation types, which has long established its
rich species diversity, and more species of Amolops may be found
through more field surveys in this nature reserve.
Consent for publication
All authors agree for publication of this article.
Declaration of competing interest
The authors declare that they have no known competing
financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
CRediT authorship contribution statement
Shuo Liu: Writing ereview and editing, Writing eoriginal
draft, Methodology, Investigation, Formal analysis, Conceptualiza-
tion. Mian Hou: Writing ereview and editing, Methodology,
Conceptualization. Mingzhong Mo: Validation, supervision,
Investigation, Conceptualization. Yi Lu: Validation, Resources,
Investigation. Jimin Guo: Validation, Resources, Investigation. Wen
Wang: Validation, Resources, Investigation. Wenxiang Zhang:
Validation, Resources. Dingqi Rao: Writing ereview and editing,
Methodology, Formal analysis, Conceptualization. Song Li: Writing
ereview and editing, Project administration, Investigation, Fund-
ing acquisition, Conceptualization.
Acknowledgments
We thank the forest rangers of Yunnan Huanglianshan National
Nature Reserve for their assistance in the fieldwork. We would like
to thank the editors and reviewers for their valuable comments on
the manuscript. This work was supported by the Project of Huan-
glian Mountains National Nature Reserve Animal Diversity Expe-
dition (grant no. E2023HLS001), National Natural Science
Foundation Project: Classificatory and Phylogenetic Studies on the
Amolops frogs of China (grant no. NSFC-31772424), and the project
from the Ministry of Ecology and Environment of China: Investi-
gation and assessment of amphibians and reptiles in southern
Yunnan.
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Please cite this article as: Liu S et al., A new species of the Amolops monticola group (Anura,Ranidae) from southern Yunnan, China, Journal of
Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2024.08.002