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Zoological Letters
A new species ofcascade frog (Anura:
Ranidae: Amolops) fromcentral Yunnan, China
Shangjing Tang1,2†, Tao Sun1,2†, Shuo Liu3†, Sangdi Luo1,2, Guohua Yu1,2* and Lina Du1,2*
Abstract
A new species of the genus Amolops, Amolops ailao sp. nov., is described from central Yunnan, China. The new species
belongs to the A. mantzorum species group. Phylogenetic analyses based on the combination of mitochondrial 16S
rRNA, COI, and cytb genes revealed that the new species is the sister taxon to Amolops ottorum with strong support.
Genetically, the new species differs from A. ottorum by 5.0% in cytb sequences. Morphologically, the new species
can be distinguished from known congeners by the combination of the following characters: true dorsolateral folds
absent, but dorsolateral folds formed by series of glands present; circummarginal groove on tip of first finger absent;
body size small (males SVL 33.0–35.1 mm and female SVL 41.3 mm); HW/SVL 0.32‒0.35; UEW/SVL 0.08‒0.10; THL/
SVL 0.52‒0.56; vomerine teeth absent; interorbital distance narrower than internarial distance; tympanum distinct,
less than half eye diameter; supratympanic fold present, indistinct; a pair of large tubercles on sides of cloaca; tibiotar-
sal articulation reaching beyond anterior corner of eye; and vocal sac absent. The cladogenesis events within the A.
mantzorum group rapidly occurred from Pliocene 4.23 Mya to Pleistocene 1.2 Mya, coinciding with the recent inten-
sive uplift of the Qinghai-Tibetan Plateau since the Pliocene. Combining findings in this study with the most recent
taxonomic progress, we consider that there are 20 known Amolops species in Yunnan, China, accounting for the high-
est proportion of amphibian diversity of Yunnan, and five of them belong to the A. mantzorum group. Among differ-
ent subfauna and water systems in Yunnan, the species diversity of Amolops in northwestern Yunnan and Nu River
Basin is highest.
Keywords Amolops mantzourm, Species group, The Qinghai-Tibetan Plateau, 16S rRNA, COI, Cytb
Introduction
e cascade frogs of genus Amolops Cope, 1865 [1]
inhabit rocky streams or waterfalls, enabled by abdom-
inal suckers in larvae and enlarged digital discs in
adults [2], and are widely distributed from Nepal and
northern India eastwards to China and southwards to
Malaysia [3]. e species diversity in Amolops has been
poorly understood owing to morphological conserva-
tion [4, 5], and efforts relying on molecular data during
the last decade have greatly improved our understand-
ing of the taxonomy and species diversity of this genus,
with a high number of new species having been dis-
covered (e.g., [3, 4, 6–13]). So far, as the most speciose
genus within the family Ranidae, the genus Amolops
contains 79 species [14], which can be allocated 10
†Shangjing Tang, Tao Sun and Shuo Liu contributedequally to this work.
*Correspondence:
Guohua Yu
yugh2018@126.com
Lina Du
dulina@mailbox.gxnu.edu.cn
1 Key Laboratory of Ecology of Rare and Endangered Species
and Environmental Protection, Guangxi Normal University, Ministry
of Education, Guilin 541004, China
2 Guangxi Key Laboratory of Rare and Endangered Animal Ecology,
College of Life Science, Guangxi Normal University, Guilin 541004, China
3 Kunming Natural History Museum of Zoology, Kunming Institute
of Zoology, Chinese Academy of Sciences, Kunming 650223, China
Page 2 of 19
Tangetal. Zoological Letters (2023) 9:15
species groups [11]. In China, a total of 50 Amolops
species have been recorded [15] and most of them have
been assigned to eight species groups, namely Amolops
chayuensis group, Amolops daiyunensis group, Amolops
hainanensis group, Amolops mantzorum group,
Amolops monticola group, Amolops marmoratus group,
Amolops viridimaculatus group, and Amolops ricketti
group, based on morphological and molecular evidence
[4, 7, 9, 11, 16–24].
e A. mantzorum species group was defined based
on the absence of true dorsolateral folds (not formed
by incomplete series of glands), circummarginal groove
on the tip of first finger, tarsal fold and tarsal glands
absent, and nuptial pad present on first finger in males
[11, 16, 17, 25]. It was comprised of 11 species [11,
24], namely Amolops mantzorum (David, 1872) [26],
Amolops granulosus (Liu and Hu, 1961) [27], Amolops
loloensis (Liu, 1950) [28], Amolops lifanensis (Liu, 1945)
[29], Amolops xinduqiao Fei, Ye, Wang, and Jiang, 2017
[25], Amolops jinjiangensis Su, Yang, and Li, 1986 [30],
Amolops tuberodepressus Liu and Yang, 2000 [31],
Amolops sangzhiensis Qian, Xiang, Jiang, Yang, and
Gui, 2023 [24], Amolops shuichengicus Lyu and Wang,
2019 [20], Amolops ottorum Pham, Sung, Pham, Le,
Zieger, and Nguyen, 2019 [3], and Amolops minutus
Orlov and Ho, 2007 [32]. Recently, A. xinduqiao was
placed into synonymy of A. mantzorum as a subspecies
[33]. us, currently the A. mantzorum species group
contains 10 species, of which two (A. ottorum and A.
minutus) are only known from northwestern Vietnam
and seven are known from southwestern China [14].
Yunnan is located in southwestern China and har-
bors a rich amphibian fauna in terms of species count
and endemism. It has been known that there are four
members of the A. mantzorum group in Yunnan, i.e., A.
jinjiangensis, A. loloensis, A. mantzorum, and A. tuber-
odepressus [15]. In recent years, a series of new species
or new records of Amolops have been discovered inten-
sively from southwestern China [2, 5, 7, 11, 18, 20, 22–
25, 34–36], suggesting that species diversity of Amolops
in the region still remains underestimated and prob-
ably more species would be found. During recent field
surveys in central Yunnan, China, we collected seven
specimens of an Amolops species that morphologically
resemble some members of the A. mantzorum group
in that they lack a circummarginal groove on tip of the
first finger and have folds formed by incomplete series
of glands along the dorsolateral junction of the body
(hereafter dorsolateral glandular folds). Molecular and
morphological comparison supported that these speci-
mens differ from other members of the genus Amolops.
us, we considered them to represent a new Amolops
species.
Materials andmethods
Sampling
Specimens were collected at Mt. Ailao, Xinping County,
Yunnan Province, China (Fig.1) by Guohua Yu in May
2019 and July 2019, and by Shuo Liu in June 2022. Speci-
mens were photographed, euthanized, fixed, and then
stored in 75% ethanol. Liver tissues were preserved in
99% ethanol. Specimens were deposited at Guangxi Nor-
mal University (GXNU) and Kunming Institute of Zool-
ogy, Chinese Academy of Sciences (KIZ).
Morphology
Morphometric data were taken using digital calipers to
the nearest 0.1mm. Morphological terminologies follow
Fei etal. [25]. Measurements included: snout-vent length
(SVL, from tip of snout to vent); head length (HL, from
tip of snout to rear of jaws); head width (HW, width of
head at its widest point); snout length (SL, from tip of
snout to anterior border of eye); internarial distance
(IND, distance between nares); interorbital distance
(IOD, minimum distance between upper eyelids); upper
eyelid width (UEW, maximum width of upper eyelid); eye
diameter (ED, diameter of exposed portion of eyeball);
tympanum diameter (TD, the greater of tympanum verti-
cal and horizontal diameters); forearm and hand length
(FHL, from elbow to tip of third finger); thigh length
(THL, from vent to knee); tibia length (TL, from knee to
heel); foot length (FL, from proximal end of inner met-
atarsal tubercle to tip of fourth toe); length of foot and
tarsus (TFL, from tibiotarsal joint to tip of fourth toe);
and horizontal diameter of digital disc of third finger
(F3DSC). Comparative morphological data of congeners
were taken from their original descriptions or re-descrip-
tions [2, 3, 6–13, 17–20, 22–25, 27–32, 34, 35, 37–59].
A multivariate principal component analysis (PCA)
was conducted using SPSS 17.0 (SPSS Inc., USA) based
on a correlation matrix of size-standardized measure-
ments (all measurements divided by SVL). Scatter
plots of the first two PCA factors were used to exam-
ine the differentiation between the new species and A.
ottorum, which was recovered as the sister to the new
species by phylogenetic analyses (see below). e meas-
urements of A. ottorum were obtained from its original
description [3].
Molecular analyses
Total genomic DNA was extracted from liver tissues. Tis-
sue samples were digested using proteinase K, and subse-
quently purified following a standard phenol/chloroform
isolation and ethanol precipitation. Fragments encod-
ing partial 16S rRNA (16S), partial cytochrome oxidase
subunit I (COI), and partial cytochrome b (cytb) genes
were amplified using primer pairs L2188 [60] /16H1 [61],
Page 3 of 19
Tangetal. Zoological Letters (2023) 9:15
Chmf4/Chmr4 [62], and F1/R3 [63], respectively. PCR
amplifications were performed in 50µl reactions using the
following cycling conditions: an initial denaturing step at
95°C for 4min; 35 cycles of denaturing at 94°C for 60s,
annealing at 46, 49, or 51°C for 60s (46°C for COI, 49°C
for cytb, and 51°C for 16S), and extending at 72°C for 60s;
and a final extending step of 72°C for 10min. Sequenc-
ing was conducted directly using the corresponding PCR
primers. All new sequences have been deposited in Gen-
Bank under Accession Nos. MN650737–MN650749,
MN650751‒MN650757, OP879227, OP880242, and
OP887035 (Table1). Homologous sequences of 25 con-
geners were obtained from GenBank. Two Odorrana spe-
cies were included as outgroups according to Ngo etal.
[64] and their sequences were also downloaded from
GenBank.
Sequences were aligned using MUSCLE with the
default parameters in MEGA 7 [65]. Uncorrected pair-
wise distances between species were calculated in MEGA
7. Because sequences of the three genes are not all avail-
able for each species (Table1), we prepared a combined
dataset of the three genes for phylogenetic analyses. e
best substitution model was selected using the Corrected
Akaike Information Criterion (AICc) in jMODELTEST
v. 2.1.6 [66] via Cipres Science Gateway [67]. Bayesian
inference was performed in MRBAYES 3.2.6 [68] based
on the selected substitution model (GTR + I + G). Two
runs were performed simultaneously with four Markov
chains starting from random tree. e chains were run
for 3,000,000 generations and sampled every 100 gen-
erations. Burn-in was checked using the program Tracer
v.1.6 [69]. e first 25% of the sampled trees were dis-
carded as burn-in and then the remaining trees were
used to create a consensus tree and to estimate Bayesian
posterior probabilities (BPPs). In addition, a maximum
likelihood (ML) analysis was conducted in RAXML-HPC
v.8.2.12 [70] with 1000 rapid bootstrap replicates.
We estimated the lineage divergence times using an
uncorrelated lognormal relaxed molecular clock model
in BEAST v. 1.8.0 [71]. e birth-death process was cho-
sen as the tree prior because of the mixed inter- and
intraspecies sampling in the dataset sets [72]. Recently,
the crown age for Amolops was inferred to be 25.01Ma
(95% HPD: 21.31‒28.58) by Wu etal. [4]. erefore, the
time of most recent common ancestor (TMRCA) of the
genus Amolops (25.01 ± 2.2Ma) was used as a second-
ary calibration point based on Wu et al. [4]. A run of
20million generations was conducted by sampling every
1000 generations. e effective sample sizes of values
of parameters were examined using Tracer v.1.6 [73].
Fig. 1 Map showing the collection site of Amolops ailao sp. nov. from central Yunnan, China (red star) and type localities of other known species
and subspecies of A. mantzorum group (black circles)
Page 4 of 19
Tangetal. Zoological Letters (2023) 9:15
Table 1 Species used for molecular analyses in this study
Species Voucher Locality 16S COI CYTB
Odorrana wuchuanensis LBML 5230 Libo, Guizhou, China KU680791 KU680791 KU680791
Odorrana margaretae HNNU1207003 - KJ815050 KJ815050 KJ815050
Amolops afghanus SYS a003852 Tongbiguan, Yunnan, China MG991895 MG991924 -
Amolops bellulus - - DQ204473 KU243079 -
Amolops chayuensis SYS a007509 Baxoi, Xizang, China MK573820 MK568333 -
Amolops chaoqin XM5526 Wenxian, Gansu, China KX645666 KX645666 KX645666
Amolops cremnobatus ROM 14528 Khe Moi, Nghe An, Vietnam DQ204477 - -
Amolops daiyunensis SYS a001739 Mt. Daiyun, Fujian, China MK263243 KX507328 -
Amolops granulosus SYS a005316 Mt. Wawu, Sichuan, China MK604851 MK605609 -
Amolops granulosus 20130258 Mt. Wawu, Sichuan, China MH922934 MH922934 MH922934
Amolops granulosus 0700332 Anxian, Sichuan, China - - KJ008439
Amolops hainanensis SYS a005281 Mt. Wuzhi, Hainan, China MK263281 MG991916 -
Amolops hongkongensis DYT W-WYS-001 Mt. Wuyi, Fujian, China KX233864 KX233864 KX233864
Amolops jinjiangensis SYS a004571 Mt. Gaoligong, Yunnan, China MK573801 MK568316 -
Amolops jinjiangeniss SCUM050435CHX Deqing, Yunnan, China EF453741 MN961403 -
Amolops jinjiangeniss CIB-XM6120 Benzilan, Deqing, Yunnan, China MZ292455 MZ292455 MZ292455
Amoops jinjiangensis KIZ047095 Chuxiong, Yunnan, China MN953701 MN961404 -
Amolops “jinjiangensis” IOZ4373 Zhongdian, Yunnan, China - - KJ008379
Amolops loloensis SYS a005351 Zhaojue, Sichuan, China MK573806 MK568321 -
Amolops loloensis SM-ZDTW-01 Shimian, Sichuan, China KT750963 KT750963 KT750963
Amolops loloensis 0700211 Huanyuan, Sichuan, China - - KJ008427
Amolops tuberodepressus CIB-XM3125 Jingdong, Yunnan, China KR559270 KR559270 KR559270
Amolops tuberodepressus YU20160272 Mt. Ailao, Xinping, Yunnan, China MN650757 - MN650749
Amolops tuberodepressus SYS a003931 Mt. Wuliang, Jingdong, Yunnan, China MG991904 MG991933 -
Amolops lifanensis SYS a005378 Lixian, Sichuan, China MK604870 MK605628 -
Amolops lifanensis 0700039 Maoxian, Sichuan, China - - KJ008445
Amolops medogensis SYS a006657 Medog, Xizang, China MK573813 MK568328 -
Amolops mantzorum mantzorum - Mt. Xiling Snow, Dayi, Sichuan, China KJ546429 KJ546429 KJ546429
Amolops mantzorum mantzorum SYS a005362 Fengtongzhai NR, Baoxing, Sichuan, China MG991893 MG991922 -
Amolops mantzorum mantzorum 0700114 Longdong, Baoxing, Sichuan, China - - KJ008297
Amolops mantzorum mantzorum 0700229 Chongzhou, Sichuan, China - - KJ008339
Amolops mantzorum ssp. 0700040 Maoxian, Sichuan, China - - KJ008277
Amolops mantzorum ssp. 0700267 Wenxian, Gansu, China - - KJ008360
Amolops mantzorum xinduqiao LCLH017 Luhuo, Sichuan, China - - KJ008392
Amolops mantzorum xinduqiao 0700307 Yajiang, Sichuan, China - - KJ008410
Amolops mantzorum xinduqiao CIB-999 Kangding, Sichuan, China - - KJ008415
Amolops marmoratus CAS240593 Mon, Myanmar JF794456 - -
Amolops marmoratus KUHE19089 Chiang Mai, Thailand AB211486 - AB259738
Amolops ottorum IEBR 4342 Muong La, Son La, Vietnam - - MK941135
Amolops ottorum TBU 06 Muong La, Son La, Vietnam - - MK941136
Amolops viridimaculatus SYS a003813 Mt. Gaoligong, Yunnan, China MK604836 MK605597 KJ008459
Amolops torrentis SYS a005289 Mt. Wuzhi, Hainan, China MK263284 MG991930 -
Amolops sangzhiensis CSUFT905 Mt. Doupeng, Hunan, China OQ079539 OQ078904 -
Amolops sangzhiensis CSUFT907 Mt. Doupeng, Hunan, China OQ079540 OQ078905 -
Amolops shuichengicus SYS a004956 Shuicheng, Guizhou, China MK604845 MK605603 -
Amolops shuichengicus SYS a004957 Shuicheng, Guizhou, China MK604846 MK605604 -
Amolops spinapectoralis ROM 7513 Tram Lap, Gia Lai, Vietnam DQ204487 - -
Amolops ricketti WUSTW01 Mt. Wugong, Jiangxi, China KF956111 KF956111 KF956111
Amolops wuyiensis - Mt. Wuyi, Fujian, China KJ933509 KJ933509 KJ933509
Amolops ailao sp. nov. GXNU YU000001 Mt. Ailao, Xinping, Yunnan, China MN650751 MN650737 MN650743
Page 5 of 19
Tangetal. Zoological Letters (2023) 9:15
e maximum clade credibility tree was constructed in
TreeAnnotator v.1.8.0 [71] and was visualized in FigTree
v.1.4.0 (from http:// tree. bio. ed. ac. uk/ softw are/ figtr ee).
Results
Phylogeny anddivergence time estimation
Alignments of 16S, COI, and cytb genes were 884 bp,
676bp, and 998bp in length, respectively. e newly col-
lected individuals from Mt. Ailao, Yunnan, China repre-
sented a distinct lineage nested in the clade of members
of the A. mantzorum group, and it was recovered as the
sister taxon to A. ottorum with strong supports (Fig.2).
Genetic distances (p-distance) between this lineage and
other species in the A. mantzorum group ranged from
1.9% (vs. A. sangzhiensis) to 8.3% (vs. A. lifanensis) in
16S (Table 2), from 4.0% (vs. A. mantzorum mantzo-
rum) to 14.2% (vs. A. lifanensis) in COI (Table2), and
from 5.0% (vs. A. ottorum) to 14.4% (vs. A. lifanensis) in
cytb (Table3). e specimen IOZ4373, which was identi-
fied as A. jinjiangensis in Lu etal. [63], was not clustered
together with topotypes of A. jinjiangensis (Fig.2).
e dating analysis revealed that A. lifanensis split from
all other members of the A. mantzorum species group ca.
7.59 Mya (95% HPD: 4.6‒10.67 Mya) and then the line-
age divergence within the A. mantzorum species group
mainly occurred from 4.23 Mya (95% HPD: 2.56‒6.1
Mya) to 1.2 Mya (95% HPD: 0.53‒1.98 Mya) (Fig.3). e
divergence between the novel lineage from Mt. Ailao and
A. ottorum occurred 2.2 Mya (95% HPD: 1.1‒3.37 Mya).
Morphometric analysis
Morphological measurements are given in Table4. We
retained the first two principal components, which had
eigenvalues above 1.0 and accounted for 64.035% of total
variance (Table5). Loadings for PC1, which accounted
for 34.26% of the total variance, were most heavily loaded
on HW, UEW, ED, and THL (load factor > 0.7), and dif-
ferentiation was found along the PC1 axis between the
new species and A. ottorum (Fig.4), indicating that the
new species differs from A. ottorum by wider head, wider
upper eyelid, larger eye, and longer femur. e second
principal component (PC2) accounted for 29.78% of
the total variance, but no clear separation was observed
along this axis between the new species and A. ottorum.
In addition, the new lineage can be distinguishable from
its congeners by body size and the combination of texture
and coloration pattern.
Taxonomic account
Amolops ailaosp. nov. (Figs.5, 6, 7 and 8; Table4)
Zoobank
Urn:lsid:zoobank.org:act:725D8480-7921-46EF-975B-
524EA75EF1A4.
Holotype
GXNU YU000004, an adult male, collected on 12 May
2019 by Guohua Yu from Mt. Ailao (23°56′58.20″N,
101°29′51.66″E, 2043m above sea level; Fig.1), Xinping
County, Yunnan Province, China.
Paratypes
GXNU YU000001–YU000003, three adult males, col-
lected at same time as the holotype from the type local-
ity by Guohua Yu; GXNU YU20160273 and GXNU
YU20160274, two adult males, collected from the type
locality by Guohua Yu on 17 July 2017; and KIZ 2022041,
an adult female, collected from the type locality by Shuo
Liu on 22 June 2022.
Etymology
The specific epithet is named for the type locality,
Ailao Mt., Xinping County, Yunnan Province, China.
We suggest the English common name “Ailao cascade
frog” and the Chinese common name “Āi Láo Tūan Wā
(哀牢湍蛙) ”.
Diagnosis
Morphologically, Amolops ailao sp. nov. resembles mem-
bers of the A. mantzorum group in the absence of true
dorsolateral folds and circummarginal groove on the disc
of the first finger, and further resembles A. jinjiangensis
and A. shuichengicus in the presence of folds formed by
incomplete series of glands along the dorsolateral junc-
tion of the body (dorsolateral glandular folds). Phylo-
genetically, a clade consisting of the new species, A.
mantzorum, A. sangzhiensis, A. jinjiangensis, A. granu-
losus, A. loloensis, A. tuberodepressus, A. shuichengicus,
Table 1 (continued)
Amolops ailao sp. nov. GXNU YU000002 Mt. Ailao, Xinping, Yunnan, China MN650752 MN650738 MN650744
Amolops ailao sp. nov. GXNU YU000003 Mt. Ailao, Xinping, Yunnan, China MN650753 MN650739 MN650745
Amolops ailao sp. nov. GXNU YU000004 Mt. Ailao, Xinping, Yunnan, China MN650754 MN650740 MN650746
Amolops ailao sp. nov. GXNU YU20160273 Mt. Ailao, Xinping, Yunnan, China MN650755 MN650741 MN650747
Amolops ailao sp. nov. GXNU YU20160274 Mt. Ailao, Xinping, Yunnan, China MN650756 MN650742 MN650748
Amolops ailao sp. nov. KIZ 2022041 Mt. Ailao, Xinping, Yunnan, China OP879227 OP880242 OP887035
Page 6 of 19
Tangetal. Zoological Letters (2023) 9:15
and A. ottorum was strongly supported (Clade I; Fig.2).
Amolops ailao sp. nov. can be distinguished from its
congeners by the combination of the following charac-
ters: (1) body size small (SVL 33.0–35.1 mm in males
and 41.3mm in female); (2) HW/SVL 0.32‒0.35; UEW/
SVL 0.08‒0.10; THL/SVL 0.52‒0.56; (3) vomerine teeth
absent; (4) tympanum distinct, less than half eye diam-
eter; (5) supratympanic fold present, indistinct; (6) true
dorsolateral folds absent, but dorsolateral glandular folds
distinct; (7) absence of circummarginal groove on the
disc of the first finger; (8) tibiotarsal articulation reaching
beyond anterior corner of eye; (9) dorsal surface smooth
with no white spines; (10) a pair of large tubercles on
sides of cloaca; 11) vocal sac absent; 12) toes fully webbed
except the fourth; 13) interorbital space narrower than
internarial space.
Description ofholotype
Adult male (SVL 33.6 mm); head slightly longer (HL
11.0 mm) than wide (HW 10.8 mm); snout obtusely
pointed, projecting beyond margin of lower jaw in ven-
tral view, rounded in profile; canthus rostralis distinct,
curved; loreal region sloping, concave; nostril oval, lat-
eral, slightly protuberant; internarial distance (IND
3.7mm) greater than interorbital distance (IOD 3.3mm);
upper eyelid width (UEW 3.0mm) slightly narrower than
interorbital distance; pineal spot present; pupil oval, hor-
izontal; tympanum distinct, rounded, less than half eye
diameter; supratympanic fold indistinct; vomerine teeth
Fig. 2 Bayesian phylogram of Amolops inferred from the combination of 16S rRNA, COI and cytb sequences. Numbers above and below branches
are Bayesian posterior probabilities and ML bootstrap values (only values above 50% are shown), respectively
Page 7 of 19
Tangetal. Zoological Letters (2023) 9:15
absent; choanae oval; tongue attached anteriorly, cordi-
form, notched posteriorly; vocal sac opening absent.
Forelimbs robust, relative length of fingers
I < II < IV < III; tips of outer three fingers expanded into
discs with circummarginal grooves, relative size of discs:
1 < 2 < 3 = 4; nuptial pads present on finger I; webbing
between fingers absent; subarticular tubercles prominent
and rounded, formula 1, 1, 2, 2; supernumerary tubercles
present; thenar (inner metacarpal) tubercle oval; outer
metacarpal tubercle single, rounded.
Hindlimbs long, heels overlapping when legs at right
angle to body, tibiotarsal articulation reaching beyond
anterior corner of eye; tibia length (TL 20.1mm) longer
than forearm and hand length (FHL17.6 mm), thigh
length (THL 18.4 mm), and foot length (FL 19.8mm);
relative length of toes I < II < I II < V < IV; all toe tips
expanded into discs with circummarginal grooves; web-
bing between toes well developed, two third web, web-
bing formula I1–1.5II1–1.5III1–2IV2–1V; subarticular
tubercles distinct, formula 1, 1, 2, 3, 2; inner metatar-
sal tubercle prominent, oval; outer metatarsal tubercle
absent; supernumerary tubercles absent.
True dorsolateral folds absent, but folds formed by
incomplete series of glands along dorsolateral junc-
tion of body (dorsolateral glandular folds) present,
extending from rear of eye to groin; skin smooth, with
a few flattened tubercles on flanks and dorsal surface
of limbs; a few small tubercles on posterior surface of
thigh and around vent; a pair of relatively large tuber-
cles at the side of the anus; ventral surface smooth; a
rictal gland.
Table 2 Genetic distance (%) between members of the A. mantzorum group estimated from 16S (lower triangle) and COI sequences
(upper triangle). A. m. mantzorum = A. mantzorum mantzorum; A. m. ssp. = A. mantzorum ssp.; A. m. xinduqiao = A. mantzorum xinduqiao
12345678910111213
1 A. ailao sp. nov. ‒NA 4.7 5.8 4.0 NA NA NA 5.5 5.4 4.7 14.2 4.5
2 A. ottorum NA ‒NA NA NA NA NA NA NA NA NA NA NA
3 A. loloensis 2.1 NA ‒3.1 3.5 NA NA NA 5.3 5.1 4.6 12.6 3.0
4 A. jinjiangensis 2.1 NA 1.0 ‒4.6 NA NA NA 5.7 5.2 4.9 13.7 4.7
5 A. m. mantzorum 2.2 NA 2.0 2.1 ‒NA NA NA 5.1 4.9 4.6 12.8 3.9
6 A. m. ssp. NA NA NA NA NA ‒NA NA NA NA NA NA NA
7 A. “jinjiangensis” NA NA NA NA NA NA ‒NA NA NA NA NA NA
8 A. m. xinduqiao NA NA NA NA NA NA NA ‒NA NA NA NA NA
9 A. tuberodepressus 2.7 NA 1.7 1.8 2.1 NA NA NA ‒6.0 5.2 13.8 5.9
10 A. granulosus 2.7 NA 2.0 2.2 2.7 NA NA NA 2.2 ‒4.1 13.0 5.8
11 A. shuichengicus 3.0 NA 2.1 2.0 3.4 NA NA NA 3.0 2.9 ‒12.4 5.5
12 A. lifanensis 8.3 NA 7.6 7.5 9.0 NA NA NA 8.3 7.8 8.3 ‒12.4
13 A. sangzhiensis 1.9 NA 1.0 0.8 2.0 NA NA NA 1.9 2.1 2.0 8.0
Table 3 Genetic distance (%) between members of the A. mantzorum group estimated from cytb sequences. A. m. mantzorum = A.
mantzorum mantzorum; A. m. ssp. = A. mantzorum ssp.; A. m. xinduqiao = A. mantzorum xinduqiao
1234567891011
1 A. ailao sp. nov. ‒
2 A. ottorum 5.0 ‒
3 A. jinjiangensis 6.4 5.1 ‒
4 A. m. xinduqiao 6.7 6.5 6.1 ‒
5 A. tuberodepressus 6.7 5.9 5.8 6.8 ‒
6 A. m. ssp. 7.0 7.0 6.6 2.4 7.1 ‒
7 A. m. mantzorum 7.0 7.6 6.7 2.5 7.1 2.9 ‒
8 A. “jinjiangensis” 7.3 7.0 6.9 3.4 6.7 3.1 3.2 ‒
9 A. loloensis 7.0 5.9 5.1 5.8 6.5 7.1 6.8 7.0 ‒
10 A. granulosus 7.8 6.7 5.6 6.7 7.1 6.9 6.6 6.9 6.0 ‒
11 A. lifanensis 14.4 14.8 14.6 14.2 14.8 14.5 13.8 15.0 14.4 13.6 ‒
12 A. shuichengicus NA NA NA NA NA NA NA NA NA NA NA
Page 8 of 19
Tangetal. Zoological Letters (2023) 9:15
Color ofholotype
In life, iris light brown with dark wash; top of head and
dorsum golden brown with large rounded black brown
and green spots; sides of head with a pale green stripe
extending from loreal region to region behind and below
eye along upper lip; a short brown stripe below the green
stripe on the loreal region; a black brown band from the
tip of the snout through the nostril to an anterior bor-
der of the eye, continuing behind the eye to the shoul-
der; temporal region black brown with a green blotch;
flanks green with few back brown and light yellow spots,
a golden brown patch below dorsolateral glands; rictal
gland pale green; limbs dorsally golden brown with black
brown bands; anterior and posterior of forelimb and
thigh black brown, mottled with green and light green
blotches; throat, chest, and venter creamy white, mot-
tled with light green and marbled with gray on throat and
chest; venter of limbs flesh-colored, scattered with light
green spots; web orange yellow.
In preservative, color faded. Dorsal surface light brown
with beige brown and gray blue spots on head and body
and beige brown bands on limbs; ventral surface white,
marbled with brown on throat and chest.
Sexual dimorphism
Body size of males smaller than that of female; nuptial
pads present on the base of finger Iin males.
Morphological variation
Color of dorsal surfaces varied among specimens.
Ground color of dorsal surfaces of the holotype and
three paratypes (GXNU YU000002, GXNU YU000003,
and KIZ2022041) is brown, and ground color of dorsal
surfaces of remaining paratypes (GXNU YU20160273,
Fig. 3 Divergence time estimates within Amolops using BEAST. Numbers above branches are average ages and blue bars represent 95% intervals.
The calibration point is highlighted with a circle
Page 9 of 19
Tangetal. Zoological Letters (2023) 9:15
GXNU YU20160274, and GXNU YU000001) is green.
No large black brown spot on dorsum and flanks of
GXNU YU20160273.
Distribution andecology
e new species is only known from the type local-
ity. It was found on leaves or small branches less than
2m above the ground along a mountain stream at night
(Fig.9) from May to July. All male types have nuptial pads
on first finger and the female paratype (KIZ 2022041) is
pregnant with eggs, suggesting that the breeding season
may be from May to July. No tadpoles were collected
for the new species. Amolops tuberodepressus was also
encountered during surveys at the type locality.
Comparison
Within the A. mantzorum group, the new species can be
distinguished from its sister taxon, A. ottorum, by smaller
body size (female SVL 41.3 mm vs. 47.5‒48.2 mm in
females), wider head (HW/SVL 0.32‒0.35 in the new spe-
cies vs. HW/SVL 0.31 in A. ottorum), wider upper eye-
lid (UEW/SVL 0.08‒0.10 in the new species vs. 0.07 in
A. ottorum), larger eye (ED/SVL 0.12‒0.14 vs. 0.12), and
longer femur (THL/SVL 0.52‒0.56 vs. 0.49), the presence
of dorsolateral glandular folds (versus absent), the pres-
ence of tubercles on flanks and limbs (versus absent), and
the presence of a pair of large tubercles on sides of cloaca
(versus absent). Amolops tuberodepressus also occurred
at the type locality of the new species. Amolops ailao sp.
nov. can be easily distinguished from A. tuberodepressus
by smaller body size (males SVL 33.0–35.1 and female
SVL 41.3mm vs. males SVL 44.3–56.7 and females SVL
60.8–71.1mm), vomerine teeth absent (vs. present), dor-
solateral glandular folds present (versus absent), a pair of
large tubercles on sides of cloaca (vs. absent), and tibio-
tarsal articulation reaching beyond anterior corner of eye
(vs. tibiotarsal articulation reaching beyond tip of snout).
e new species is distinguishable from A. minu-
tus by vomerine teeth absent (vs. strongly developed),
vocal sac absent (vs. paired well-developed vocal sacs),
and tympanum less than half eye diameter (vs. TD/
Table 4 Measurements (mm) of the holotype and paratypes of Amolops ailao sp. nov. Abbreviations defined in the text
Character GXNU
YU000001 GXNUYU000002 GXNUYU000003 GXNUYU000004 GXNUYU20160273 GXNUYU20160274 KIZ 2022041
Sex ♂ ♂ ♂ ♂ ♂ ♂ ♀
SVL 33.0 35.1 33.4 33.6 33.7 33.4 41.3
HL 11.2 11.6 11.3 11.0 11.1 10.7 14.7
HW 10.8 11.4 11.1 10.8 11.0 10.6 14.4
SL 4.6 4.9 4.8 4.8 4.5 4.9 6.2
IND 3.5 3.9 3.7 3.7 3.7 3.6 5.0
IOD 3.2 3.3 3.1 3.3 3.1 3.1 3.4
UEW 3.2 2.8 3.3 3.0 3.3 3.0 3.6
ED 4.5 4.4 4.6 4.3 4.5 4.3 5.0
TD 1.6 1.6 1.6 1.7 1.6 1.4 1.9
FHL 17.5 18.5 17.1 17.6 17.2 17.3 22.0
THL 18.5 18.9 18.6 18.4 17.6 18.3 22.2
TL 19.2 20.0 19.1 20.1 18.7 19.1 23.2
TFL 26.7 29.1 26.9 29.5 26.7 27.0 32.8
FL 18.3 20.1 18.5 19.8 18.4 18.6 22.0
F3DSC 1.8 2.0 1.8 1.8 1.8 1.6 2.1
Table 5 Factor loadings of first two principal components of 12
size-adjusted morphometric characteristics of Amolops ailao sp.
nov. and A. ottorum
Character PC1 PC2
Eigenvalue 4.111 3.573
% variation 34.257% 29.778%
HW (head width) 0.808 -0.474
HL (head length) 0.566 -0.486
SL (snout length) -0.097 -0.320
IND (internarial distance) 0.258 0.774
IOD (interorbital distance) -0.469 0.787
UEW (width of upper eyelid) 0.914 -0.009
ED (eye diameter) 0.704 0.198
TD (tympanum diameter) 0.691 0.528
THL (thigh length) 0.893 0.121
TL (tibia length) 0.026 0.850
TFL (length of foot and tarsus) 0.375 0.767
F3SDC3 (diameter of disc of finger III) -0.308 0.385
Page 10 of 19
Tangetal. Zoological Letters (2023) 9:15
ED mean 0.52 in males and mean 0.58 in females). e
new species differs from the other seven members of
the A. mantzorum species group (A. sangzhiensis, A.
shuichengicus, A. mantzorum, A. granulosus, A. jin-
jiangensis, A. lifanensis, and A. loloensis) by vomerine
teeth absent (vs. present) and smaller body size, males
SVL 33.0–35.1mm and female SVL 41.3mm (vs. males
SVL 40.3–40.9mm and females SVL 52.6–57.7mm in
A. sangzhiensis, males SVL 34.6–39.6mm and females
SVL 48.5–55.5 mm in A. shuichengicus, males SVL
41.2–57.5 mm and females SVL 48.5–72.0 mm in A.
mantzorum, males SVL 36.3–41.8mm and female SVL
51.9 mm in A. granulosus, males SVL 43.0–52.0 mm
and females SVL 54–66.4mm in A. jinjiangensis, males
SVL 52–56mm and females SVL 61.0–79.0 in A. lifan-
ensis, and males SVL 54.5–62.0 mm and females SVL
69.5–77.5mm in A. loloensis).
Amolops ailao sp. nov. further differs from A. mant-
zorum, A. lifanensis, and A. loloensis by the presence of
distinct dorsolateral glandular folds (vs. absent); from A.
mantzorum, A. jinjiangensis, A. lifanensis, and A. loloen-
sis by tympanum distinct (vs. obscure or invisible); and
from A. granulosus by dorsal surface smooth with no
white spines (vs. dorsal surface rough with spines) and
vocal sac absent (vs. a pair of internal subgular vocal
sacs).
Amolops ailao sp. nov. further differs from A. jinjian-
gensis by dorsal surfaces smooth (vs. skin coarse, with
many rounded tubercles on head side, body side, and
posterior part of dorsum); from A. lifanensis by toes fully
webbed except the fourth (vs. webs fully developed to the
bases of all toe disks); and from A. loloensis by interorbi-
tal space narrower than internarial space (vs. interorbital
space about equal to the internarial space).
e A. monticola group contains 23 members,
namely A. adicola Patel, Garg, Das, Stuart, and Biju,
2021 [12], A. akhaorum Stuart, Bain, Phimmachak, and
Spence, 2010 [50], A. aniqiaoensis Dong, Rao, and Lü,
2005 [55], A. archotaphus (Inger and Chanard, 1997)
[43], A. bellulus Liu, Yang, Ferraris, and Matsui, 2000
[74], A. chakrataensis Ray, 1992 [75], A. chaochin Jiang,
Ren, Lyu, and Li, 2021 [11], A. chunganensis (Pope,
1929) [76], A. compotrix (Bain, Stuart, and Orlov, 2006)
[38], A. cucae (Bain, Stuart, and Orlov, 2006) [38], A.
daorum (Bain, Lathrop, Murphy, Orlov, and Ho, 2003)
[37], A. deng Jiang, Wang, and Che, 2020 [34], A. iri-
odes (Bain and Nguyen, 2004) [39], A. kohimaensis Biju,
Mahony, and Kamei, 2010 [40], A. mengdingensis Yu,
Wu, and Yang, 2019 [22], A. mengyangensis Wu and
Fig. 4 Scatterplot of principal components 1 and 2 of size-adjusted morphometric data for A. ailao sp. nov. and A. ottorum
Page 11 of 19
Tangetal. Zoological Letters (2023) 9:15
Tian, 1995 [52], A. monticola (Anderson, 1871) [77], A.
nyingchiensis Jiang, Wang, Xie, Jiang, and Che, 2016 [7],
A. putaoensis Gan, Qin, Lwin, Li, Quan, Liu, and Yu,
2020 [41], A. truongi Pham, Pham, Ngo, Sung, Ziegler,
and Le, 2023 [56], A. tuanjieensis Gan, Yu, and Wu,
2020 [18], A. vitreus (Bain, Stuart, and Orlov, 2006)
[38], and A. wenshanensis Yuan, Jin, Li, Stuart, and Wu,
2018 [23]. Amolops ailao sp. nov. can be distinguished
from these species by absence of true dorsolateral folds
(vs. present). e new species can be further distin-
guished from A. adicola, A. akhaorum, A. aniqiaoensis,
A. archotaphus, A. chaochin, A. chunganensis, A. com-
potrix, A. cucae, A. daorum, A. iriodes, A. kohimaen-
sis, A. mengdingensis, A. mengyangensis, A. monticola,
A. putaoensis, A. truongi, A. tuanjieensis, A. vitreus,
and A. wenshanensis by vocal sac absent (vs. present);
and from A. adicola, A. akhaorum, A. aniqiaoensis, A.
archotaphus, A. bellulus, A. chakrataensis, A. chaochin,
A. chunganensis, A. compotrix, A. cucae, A. deng, A.
iriodes, A. kohimaensis, A. mengdingensis, A. mengyan-
gensis, A. nyingchiensis, A. putaoensis, A. truongi, A.
tuanjieensis, A. vitreus, and A. wenshanensis by vomer-
ine teeth absent (vs. present).
Amolops ailao sp. nov. is distinguishable from A. chay-
uensis Sun, Luo, Sun, and Zhang, 2013 [21], the sole
member of the A. chayuensis group, by true dorsolateral
folds absent (vs. present), vocal sacs absent (vs. present),
and vomerine teeth absent (vs. present).
The A. viridimaculatus group contains 14 species
based on recent taxonomic studies [11, 13, 57], namely
A. beibengensis Jiang, Li, Zou, Yan, and Che, 2020 [33],
A. chanakya Saikia, Laskar, Dinesh, Shabnam, and
Sinha, 2022 [57], A. formosus (Günther, 1876) [78],
A. himalayanus (Boulenger, 1888) [79], A. kaulbacki
(Smith, 1940) [80], A. longimanus (Andersson, 1939)
[81], A. medogensis Li and Rao, 2005 [55], A. nidorbel-
lus Biju, Mahony, and Kamei, 2010 [40], A. pallasita-
tus Qi, Zhou, Lyu, Lu, and Li, 2019 [2], A. senchalensis
Chanda, 1987 “1986” [82], A. tawang Saikia, Laskar,
Dinesh, Shabnam, and Sinha, 2022 [57], A. wangyali
Mahony, Nidup, Streicher, Teeling, and Kamei, 2022
[13], A. wangyufani Jiang, 2020 [34], and A. viridi-
maculatus (Jiang, 1983) [45]. The new species can be
distinguished from these species by vomerine teeth
absent (vs. present), glands in compete series along
dorsolateral junction present (vs. absent), and smaller
body size (vs. male SVL 75.8 mm and females SVL
Fig. 5 Holotype of Amolops ailao sp. nov. in life (a and b) and in preservative (c and d). (a) dorsolateral view, (b) ventral view, (c) dorsal view, and (d)
ventral view
Page 12 of 19
Tangetal. Zoological Letters (2023) 9:15
90.2–93.2mm in A. beibengensis, male SVL 76.4mm
in A. chanakya, males SVL 61.3–63.1mm and females
SVL 79.4–83.7mm in A. formosus, male SVL 80mm in
A. himalayanus, males SVL 70–72mm in A. kaulbacki,
male SVL ca. 95mm and females SVL72.4–96.9mm in
A. medogensis, males SVL 76.4–82.3mm and females
SVL 85.4–98mm in A. nidorbellus, female SVL 70.6–
72.3mm in A. pallasitatus, male SVL 46.2mm in A.
senchalensis, male SVL 82.5mm in A. tawang, males
SVL 71.4–76.7mm and females SVL 80.5–89.6mm in
A. wangyali, males SVL 68.3–69.0mm and female SVL
83.4mm in A. wangyufani, males SVL 72.7–82.3 mm
and female SVL 83.0–94.3mm in A. viridimaculatus).
e Amolops marmoratus group contains 13 species,
namely A. afghanus (Günther, 1858) [83], A. assamen-
sis Sengupta, Hussain, Choudhury, Gogoi, Ahmed, and
Choudhury, 2008 [49], A. gerbillus (Annandale, 1912)
[84], A. indoburmanensis Dever, Fuiten, Konu, and
Wilkinson, 2012 [6], A. jaunsari Ray, 1992 [75], A. lato-
palmatus (Boulenger, 1882) [85], A. mahabharatensis
Khatiwada, Shu, Wang, Zhao, Xie, and Jiang, 2020 [10],
A. marmoratus (Blyth, 1855) [86], A. nepalicus Yang,
1991 [53], A. panhai Matsui and Nabhitabhata, 2006
[47], A. siju Saikia, Sinha, Shabnam, and Dinesh, 2023
[59], A. terraorchis Saikia, Sinha, Laskar, Shabnam, and
Dinesh, 2022 [58], and A. yarlungzangbo Jiang, Wang,
Li, Qi, Li, and Che, 2020 [34]. e new species differs
from these species by circummarginal groove on disc
of finger I absent (vs. present), vomerine teeth absent
(vs. present), and vocal sac absent (vs. present with the
exception of A. siju).
e new species differs from A. spinapectoralis Inger,
Orlov, and Darevsky, 1999 [41], the sole member of the A.
spinapectoralis group, by vomerine teeth absent (vs. pre-
sent), circummarginal groove on disc of finger I absent
(vs. present), and vocal sac absent (vs. present).
e A. larutensis group contains four species, namely A.
australis Chan, Abraham, Grismer, and Grismer, 2018 [8],
A. cremnobatus Inger and Kottelat, 1998 [44], A. gerutu
Chan, Abraham, Grismer, and Grismer, 2018 [8], and A.
larutensis (Boulenger, 1899) [87]. e new species can be
distinguished from these species by vomerine teeth absent
(vs. present), circummarginal groove on disc of finger I
absent (vs. present), and vocal sac absent (vs. present).
e A. ricketti group contains eight species, namely A.
shihaitaoi Wang, Li, Du, Hou, and Yu, 2022 [5], A. sin-
ensis Lyu, Wang, and Wang, 2019 [19], A. ricketti (Bou-
lenger, 1899) [88], A. wuyiensis (Liu and Hu, 1975) [46],
A. yunkaiensis Lyu, Wang, Liu, Zeng, and Wang, 2018
[9], A. albispinus Sung, Wang, and Wang, 2016 [51], A.
yatseni Lyu, Wang, and Wang, 2019 [19], and A. tonki-
nensis (Ahl, 1927 “1926’’) [89]. e new species differs
from these species by circummarginal groove on disc of
Fig. 6 Ventral view of foot of the holotype of Amolops ailao sp. nov.
in preservative
Fig. 7 Views of the female paratype of Amolops ailao sp. nov. (KIZ
2022041) in life
Page 13 of 19
Tangetal. Zoological Letters (2023) 9:15
finger I absent (vs. present), dorsolateral glandular folds
present (vs. absent), and nuptial pad without conical or
papillate nuptial spines (vs. present).
e A. daiyunensis group contains three species,
namely A. daiyunensis (Liu and Hu, 1975) [46], A. hong-
kongensis (Pope and Romer, 1951) [90], and A. teochew
Fig. 8 Dorsolateral views of male paratypes in life. (a) GXNU YU20160273; (b) GXNU YU20160274; (c) GXNU YU000003; and (d) GXNU YU000002
Fig. 9 Habitat at the type locality of Amolops ailao sp. nov. (a) and an adult male of Amolops ailao sp. nov. sitting on branches at the type locality (b)
Page 14 of 19
Tangetal. Zoological Letters (2023) 9:15
Zeng, Wang, Lyu, and Wang, 2021 [54]. e new species
differs from them by circummarginal groove on disc of
finger I absent (vs. present), dorsolateral glandular folds
present (vs. absent), and vocal sac absent (vs. present).
e A. hainanensis group contains two members,
namely A. hainanensis (Boulenger, 1900) [91] and A. tor-
rentis (Smith, 1923) [92]. e new species can be distin-
guished from them by dorsolateral glandular folds absent
(vs. absent), circummarginal groove on disc of finger
absent (vs. present), and nuptial pad present in males (vs.
absent).
e new species is distinguishable from A. bin-
chachaensis Rao, Hui, Ma, and Zhu, 2022 “2020” [35],
which has not been assigned to any species group, by true
dorsolateral folds absent (vs. present) and circumma-
rginal groove on disc of finger I absent (vs. present).
Discussion
Based on molecular and morphological evidence, we
find a novel lineage belonging to the A. mantzorum spe-
cies group from central Yunnan, China and describe it as
a new species. is finding brings the number of species
of the A. mantzorum group to 11. e phylogenetic rela-
tionships within the clade consisting of all members of
the A. mantzorum group with the exception of A. lifan-
ensis (labelled as clade I) were not well resolved and most
basal branches in this clade are short (Fig.2), suggesting
that this group might have undergone a rapid speciation
process. is inference was supported by the analysis of
divergence dating, which revealed that the cladogenesis
events within the A. mantzorum group mainly occurred
from Pliocene 4.23 Mya (95% HPD: 2.56‒6.1 Mya) to
Pleistocene 1.2 Mya (95% HPD: 0.53‒1.98 Mya) (Fig.3). It
is generally believed that the southeastern margin of the
Qinghai-Tibet Plateau has experienced rapid and recent
uplift since the Pliocene [93–95] and there were two
phases of recent intense uplift occurring between 0.6 and
3.4 Mya [93], which caused dramatic habitat and climatic
changes (e.g., reorganization of drainage [96] and for-
mation of fluvial system [97]) and created environmen-
tal conditions (new habitats, dispersal barriers, etc.) that
increase the rate at which species divide and evolve to
form new ones [98]. Additionally, so far A. minutus has
never been included in phylogenetic analysis. us, more
studies are necessary to unveil the phylogenetic relation-
ships within the A. mantzorum species group.
e phylogenetic position of A. jinjiangensis in previ-
ous studies is controversial. Lu etal. [63] found that A.
jinjiangensis is closely related to A. mantzorum, but
recently Lyu etal. [20] and Wang etal. [99] revealed that
A. jinjiangensis is the sister taxon to A. loloensis. e
samples of A. jinjiangensis in Lu et al. [63] came from
Zhongdian in Yunnan and Yajiang in Sichuan, while
samples of A. jinjiangensis in Lyu etal. [20] and Wang
etal. [99] contained topotypes from Deqin, Yunnan. In
this study, with inclusion of samples from both Zhong-
dian and the type locality (Deqin, Yunnan), we found that
A. jinjiangensis contains two clades. e clade contain-
ing topotypes was recovered as the sister to A. loloensis,
whereas the clade containing the sample from Zhongdian
was nested in the clade of A. mantzorum (Fig. 2), indi-
cating that the samples from Zhongdian in Lu etal. [63]
actually do not belong to A. jinjiangensis but belong to A.
mantzorum. Amolops mantzorum is comprised of four
sub-lineages and the central lineage refers to A. mantzo-
rum mantzorum according to Frost [14]. us, additional
studies are needed to name the northern lineage and the
lineage containing samples from Zhongdian in Yunnan.
anks to the extremely complicated topography and
climatic condition in Yunnan, which promoted rapid
divergence and speciation in small and isolated popula-
tions [100], Yunnan is the region with highest amphib-
ian species diversity in China [15]. Of the 649 amphibian
species known from China, about one-third (220 spe-
cies belonging to 51 genus) are distributed in Yunnan
(Fig. 10). Combining findings in this study with most
recent taxonomic progress [5, 13, 35, 36], we consider
that so far there are 20 Amolops species known from Yun-
nan, accounting for the highest proportion of amphibian
diversity of Yunnan (ca. 9.1%; Fig.11) and including five
members of the A. mantzorum group (namely, A. ailao
sp. nov., A. jinjiangensis, A. loloensis, A. tuberodepres-
sus, and A. mantzorum), one member of the A. marmo-
ratus group (namely A. afghanus), eight members of the
A. monticola group (namely A. bellulus, A. daorum, A.
deng, A. iriodes, A. mengdingensis, A. mengyangensis, A.
putaoensis, A. tuanjieensis, and A. wenshanensis), two
members of the A. viridimaculatus group (namely A. vir-
idimaculatus, A. kaulbacki), one member of the A. rick-
etti group (namely A. shihaitaoi), one member of the A.
chayuensis group (A. chayuensis), and A. binchachaensis,
which has not yet been assigned to any species group but
likely belongs to the A. monticola group owing to the fact
that it has true dorsolateral folds and head side dark with
white upper lip stripe. Among different parts of Yunnan,
the species diversity of Amolops in northwestern Yun-
nan was the highest (eight species, namely A. bellulus, A.
binchachaensis, A. chayuensis, A. deng, A. jinjiangensis,
A. kaubacki, A. mantzorum, A. putaoensis), followed by
western Yunnan (six species, namely A. afghanus, A. bel-
lus, A. mengdingensis, A. tuanjieensis, A. tuberodepressus,
and A. viridimaculatus), central Yunnan (four species,
namely A. daorum, A. tuberdepressus, A. viridimaculatus,
and A. ailao sp. nov.), and southeastern Yunnan (three
species, namely A. iriodes, A. shihaitaoi, and A. wenshan-
ensis) in order, while southern and northeastern Yunnan
Page 15 of 19
Tangetal. Zoological Letters (2023) 9:15
have only one species each (Fig. 12). Accordingly, spe-
cies diversity of Amolops in the Nu River Basin is high-
est (seven species, namely A. bellulus, A. binchachaensis,
A. chayuensis, A. deng, A. kaulbacki, A. tuberodepressus,
and A. viridimaculatus), followed by the Lancang River
Basin (six species, namely A. daorum, A. mengdingensis,
A. mengyangensis, A. tuanjieensis, A. tuberodepressus,
and A. viridimaculatus), the Red River Basin (five spe-
cies, namely A. iriodes, A. shihaitaoi, A. tuberodepressus,
A. wenshanensis, and A. ailao sp. nov.), the Dulong River
Basin (three species, namely A. chayuensis, A. kaulbacki,
and A. putaoensis) and the Jinsha River Basin (three spe-
cies, namely A. jinjiangensis, A. loloensis, and A. mant-
zorum), while there are two species (A. afghanus and A.
viridimaculatus) in the Basins of the Ying River and the
Ruili River (Fig.12), both of which flow to the Irrawaddy
River. Amolops ailao sp. nov. is sympatric with A. tubero-
depressus at the type locality, but it is easy to distinguish
them because the new species has smaller body size and
dorsolateral glandular folds and lacks vomerine teeth.
Amolops mantzorum was widely recorded from cen-
tral, southwestern, and northwestern Yunnan [15, 17,
101]. Certainly some of these records actually apply to
A. tuberodepressus or A. jinjiangensis because they were
once placed into the synonymy of A. mantzorum by Fei
etal. [17, 101]. Studies based on additional sampling will
be necessary to clarify the species boundary within the A.
mantzorum species group in Yunnan, China.
Conclusions
In summary, based on morphological and molecular evi-
dence, we revealed a new cascade frog species belong-
ing to the A. mantzorum species group. e new species
Amolops ailao sp. nov. is only found in Mt. Ailao, central
Yunnan, China and has considerable variation of color
pattern. Including A. ailao sp. nov., now there are 20
Amolops species known from Yunnan, China and five of
them belong to the A. mantzorum species group. We also
Fig. 10 Comparison of amphibian diversity between Yunnan and adjacent provinces on species and genus level
Fig. 11 Contributions of Amolops and other genera to the amphibian diversity in Yunnan, China
Page 16 of 19
Tangetal. Zoological Letters (2023) 9:15
revealed that the samples of A. jinjiangensis in Lu etal.
[58] are misidentification of A. mantzorum. e A. man-
tzorum species group has undergone a rapid speciation
process since the Pliocene, coinciding with the recent
rapid uplift of the Qinghai-Tibetan Plateau since the Plio-
cene. Among different subfauna and water systems in
Yunnan, northwestern Yunnan and Nu River Basin har-
bor the highest species diversity of Amolops. e findings
in this study improve our understanding of the species
diversity of the genus Amolops. More studies are neces-
sary to unveil the phylogenetic relationships and species
boundaries within the A. mantzorum species group.
Acknowledgements
We are grateful to Jian Zang and Chunsheng Du for assistance during the
surveys.
Authors’ contributions
GHY and LND conceptualized and designed the study. GHY, LND, SL, and SDL
conducted the field surveys. SJT, TS, and SL collected and analyzed morpho-
logical and molecular data, prepared figures, and drafted the manuscript. SDL
acquired images and prepared figures. All authors read and approved the final
manuscript.
Funding
This work was supported by the National Natural Science Foundation of
China (32060114, 31872212), Guangxi Natural Science Foundation Pro-
ject (2022GXNSFAA035526), and Key Laboratory of Ecology of Rare and
Endangered Species and Environmental Protection (Guangxi Normal
University), Ministry of Education (ERESEP2022Z04), and Guangxi Key Labora-
tory of Rare and Endangered Animal Ecology, Guangxi Normal University
(19-A-01-06).
Availability of data and materials
All data generated or analyzed during this study are included in this published
article. Sequences are deposited in GenBank, NCBI.
Declarations
Ethics approval and consent to participate
Not applicable.
Consent for publication
Not applicable.
Competing interests
The authors declare that they have no competing interests.
Received: 1 January 2023 Accepted: 27 June 2023
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