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A new species of Odorrana is described from the karst forests in northeastern Vietnam based on morphological differences and molecular divergence. Morphologically, the new species is distinguishable from its congeners on the basis of a combination of the following diagnostic characters: (1) size large (SVL 85.9-91.6 mm in males, 108.7-110.1 mm in females); (2) head longer than wide; (3) vomerine teeth present; (4) external vocal sacs absent; (5) snout short (SL/SVL 0.16-0.17); (6) tympanum large (TD/ED 0.70 in males, 0.68 in females); (7) dorsal surface of head and anterior part of body smooth, posterior part of body and flanks with small tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) webbing formula I0-0II0-0III0-1/2IV1/2-0V; (11) in life, dorsum green with dark brown spots; (12) flanks greyish brown with dark brown spots; (13) throat and chest grey, underside of limbs with large dark brown spots, edged in white, forming a network. In the phylogenetic analyses, the new species is unambiguously nested within the O. andersonii group, and placed as the sister taxon to O. wuchuanensis.
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Accepted by J. Rowley: 6 Jan. 2016; published: 26 Feb. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4084 (3): 421
435
http://www.mapress.com/j/zt/
Article
421
http://doi.org/10.11646/zootaxa.4084.3.7
http://zoobank.org/urn:lsid:zoobank.org:pub:9A80A330-D3A4-4F14-95EF-ECD2DF63823F
A new species of Odorrana (Amphibia: Anura: Ranidae) from Vietnam
CUONG THE PHAM
1
, TRUONG QUANG NGUYEN
1,7
, MINH DUC LE
2,3,4
,
MICHAEL BONKOWSKI
5
& THOMAS ZIEGLER
5,6
1
Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam.
E-mail: cuong03091983@yahoo.com, nqt2@yahoo.com,
2
Faculty of Environmental Sciences, Hanoi University of Science, Vietnam National University, 334 Nguyen Trai Road, Hanoi,
Vietnam. E-mail: le.duc.minh@hus.edu.vn
3
Centre for Natural Resources and Environmental Studies, Hanoi National University, 19 Le Thanh Tong, Hanoi, Vietnam
4
Department of Herpetology, American Museum of Natural History, Central Park West at 79
th
Street, New York, New York 10024
5
Zoological Institute, Department of Terrestrial Ecology, University of Cologne, Zülpicher Strasse 47b, D–50674 Cologne, Germany.
E-mail: m.bonkowski@uni–koeln.de
6
AG Zoologischer Garten Köln, Riehler Strasse 173, D–50735 Cologne, Germany. E-mail: ziegler@koelnerzoo.de
7
Corresponding author. E-mail: nqt2@yahoo.com
Abstract
A new species of Odorrana is described from the karst forests in northeastern Vietnam based on morphological differences
and molecular divergence. Morphologically, the new species is distinguishable from its congeners on the basis of a com-
bination of the following diagnostic characters: (1) size large (SVL 85.9–91.6 mm in males, 108.7–110.1 mm in females);
(2) head longer than wide; (3) vomerine teeth present; (4) external vocal sacs absent; (5) snout short (SL/SVL 0.16–0.17);
(6) tympanum large (TD/ED 0.70 in males, 0.68 in females); (7) dorsal surface of head and anterior part of body smooth,
posterior part of body and flanks with small tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10)
webbing formula I0–0II0–0III0–1/2IV1/2–0V; (11) in life, dorsum green with dark brown spots; (12) flanks greyish
brown with dark brown spots; (13) throat and chest grey, underside of limbs with large dark brown spots, edged in white,
forming a network. In the phylogenetic analyses, the new species is unambiguously nested within the O. andersonii group,
and placed as the sister taxon to O. wuchuanensis.
Key words: Odorrana mutschmanni sp. nov., karst forest, molecular phylogeny, taxonomy, Cao Bang Province
Introduction
Cascade frogs of the genus Odorrana have a wide distribution in Asia, from northeastern India and southern China
eastwards to Japan, throughout Indochina and southwards to Sumatra and Borneo (Nguyen et al. 2009; Fei et al.
2012; Mo et al. 2015; Wang et al. 2015). Odorrana belongs to one of the most diverse groups of amphibians with
currently 56 recognized species, more than 20 of which were described in the last ten years (Frost 2015). Due to
morphological similarity in the genus, species that were formally thought to be widespread are now being
recognized as complexes of species with much narrower distributions. For instance, Bain et al. (2003) described six
new species within the Odorrana livida (Rana livida) species complex, and Bain & Stuart (2005) subsequently
discovered another new species of this complex from Thailand, namely Odorrana indeprensa (Rana indeprensa)
Bain & Stuart.
During our recent field work in northeastern Vietnam, Odorrana specimens were collected in the karst forests
of Cao Bang Province. Because many species of the genus Odorrana are not easily identified owing to subtle
interspecific morphological differences (Wang et al. 2015), the specimens from Cao Bang were initially identified
as members of the Odorrana andersonii group (Clade B2 in Chen et al. 2013) based on molecular analyses. Closer
morphological examination however showed that the specimens from Cao Bang could be clearly distinguished
from other known members of the O. andersonii group by a combination of morphological features. In the
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phylogenetic analyses, the taxon from Cao Bang was distinctly separated from its congeners but embedded within
the O. andersonii species group with strong support values by three different analyses. Due to these distinctions,
we describe it herein as a new species.
Material and methods
Sampling. Field surveys were conducted in April and May 2012 by T.Q. Nguyen, H.T. An, S. Herbst, T. Lehmann,
and M. Bonkowski (hereafter T.Q. Nguyen et al.), in June 2014 by C.T. Pham, M. Bernardes, M. van Schingen
(hereafter C.T. Pham et al.), in May 2015 by T.Q. Nguyen in Ha Lang District, Cao Bang Province, northeastern
Vietnam. Specimens were collected between 19:00 and 23:00 h. After taking photographs, specimens were fixed in
90% ethanol for 8–10 h, then later transferred to 70% ethanol for permanent storage. Tissue samples were
preserved separately in 95% prior to fixation. Specimens referred to in this paper are deposited in the collections of
the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam; Vietnam National Museum of Nature
(VNMN), Hanoi, Vietnam; and the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn,
Germany.
Molecular data and phylogenetic analyses. We used Le et al.’s (2006) protocols for extraction,
amplification, and DNA sequencing. Two fragments of the mitochondrial genes 12S and 16S were amplified using
four pairs of primers, L1991 + H1478 (Kocher et al. 1989) and L33 + H56 (Chen et al. 2013) and L2A + H10
(Hedges 1994) and L3975 + H4551 (Simon et al. 1994) for 16S. In addition to newly generated sequences, we
compiled available data from closely related species from GenBank, primarily from Chen et al. (2013) (Table 1).
Two species, Babina daunchina and Odorrana chapaensis, were used as outgroups according to Chen et al. (2013).
After sequences were aligned by Clustal X v2 (Thompson et al. 1997), data were analyzed using maximum
parsimony (MP), maximum likelihood (ML), and Bayesian analysis (BA), as implemented in PAUP*4.0b10
(Swofford 2001) and MrBayes v3.2 (Ronquist et al. 2012), respectively. Settings for these analyses followed Le et
al. (2006), except that the number of generations in the Bayesian analysis was increased to 1 × 10
7
. The optimal
model for nucleotide evolution was set to GTR+I+Γ as selected by Modeltest v3.7 (Posada & Crandall 1998).
Nodal support was evaluated using Bootstrap replication (BP) as calculated in PAUP and posterior probability (PP)
in MrBayes v3.2. Gaps in the alignment were included in the analyses as they may contain important phylogenetic
information (Giribet & Wheeler 1999). Uncorrected pairwise divergences were calculated in PAUP*4.0b10.
Morphological characters. Measurements were taken with a digital caliper to the nearest 0.1 mm. The
following abbreviations were used: SVL: snout-vent length, HL: head length (from the back of mandible to tip of
snout), HW: maximum head width (across angles of jaws), SL: snout length (from anterior corner of eye to the tip
of snout), NS: distance from nostril to the tip of snout, EN: distance from anterior corner of eye to the nostril, IND:
internarial distance, IOD: interorbital distance, ED: eye diameter, UEW: maximum width of upper eyelid, DAE:
distance between anterior corners of eyes, MN: mandible to nostril, MFE back of mandible to front of the eye,
MBE back of mandible to back of the eye; DPE: distance between posterior corners of eyes, TD: tympanum
diameter, TYE: distance from anterior margin of tympanum to posterior corner of the eye, FLL: forelimb length
(from axilla to elbow), HAL: hand length (from elbow to the tip of third finger), fd1-4: width of discs of fingers I-
IV, TFL: third finger length, OPT: outer palmar tubercle length, IPT: inner palmar tubercle length, NPL: Nuptial
pad length - finger I, FeL: femur length (from vent to knee), TbL: tibia length (from knee to tarsus), TbW: tibia
width, FoL: foot length (from tarsus to the tip of fourth toe), FTL: fourth toe length, IMT: inner metatarsal tubercle
length, td1-5: width of discs of toes I-V, phalanges: width of phalanges of finger III. For webbing formula we
followed Glaw & Vences (2007).
We compared morphological characters of the new species with other members of the genus (see specimens
examined) and comparative data obtained from the literature (e.g., Bain et al. 2003; Bain & Stuart 2006 “2005”;
Bain & Stuart 2006; Bain et al. 2009; Chen et al. 2010a, b; Fei et al. 2009; Fei et al. 2010; Li et al. 2008; Liang et
al. 2001; Maeda & Matsui 1990; Matsui 1994; Matsui & Jaafar 2006; Mo et al. 2015; Orlov et al. 2003, 2006;
Stuart & Bain 2005; Stuart & Chan-ard 2005; Stuart et al. 2005, 2006; Tran et al. 2008; Wang et al. 2015; Yang
2008).
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A NEW ODORRANA FROM VIETNAM
TABLE 1. GenBank accession numbers, and associated samples that were used in this study. All sequences generated by
this study have accession numbers: KU356761–8.
Results
Phylogenetic analyses. The combined matrix contained 1836 aligned characters, 766 of 12S and 1070 of 16S. MP
analysis of the dataset recovered six most parsimonious trees with 751 steps (CI = 0.70; RI = 0.69). The first of the
six trees is shown in Fig. 1. Tree topology is generally in agreement with that of Chen et al. (2013), and the taxon
from Cao Bang is unambiguously nested within clade B2 of Chen et al. (2013) (BP = 100 and 78, PP = 97). In the
Bayesian analysis, -lnL scores reached stationarity after 10,000 generations in both runs. The Bayesian topology
was identical to the ML’s topology, and very similar to that of the MP analyses, except for the placements of
Odorrana lungshengensis and O. margaretae. The new species was supported as the sister taxon to O.
wuchuanensis in all analyses (BP = 100, PP = 100) (Fig. 1). This species is highly divergent from the latter in terms
of genetic distance with the minimum pairwise divergence of approximately 4.3%, calculated based on the
combined 12S and 16S data (Table 2).
Odorrana mutschmanni sp. nov.
(Figs. 2, 3)
Holotype: IEBR 3723 (Field No. CB 2015.12), adult male, collected by T.Q. Nguyen on 22 April 2015 in the karst
forest near Coong Village (22
o
42.712’N, 106
o
40.075’E, at an elevation of 447 m), Duc Quang Commune, Ha Lang
District, Cao Bang Province, Vietnam.
Paratypes: IEBR 3724 (Field No. CB 2012.77), adult males, collected on 15 April 2012; IEBR 3725 (Field
No. CB 2012.89), adult female and IEBR 3726–3729 (Field No. CB 2012.90–93), adult males, collected on 16
April 2012, by T.Q. Nguyen et al.; ZFMK 97329, 97330 (Field No. CB 2012.139, 2012.140), adult males, collected
on 3 May 2012, by H. T. An, S. Herbst and T. Lehmann; IEBR 3730 (Field No. CB2014.16), adult female,
collected by C.T. Pham et al. on 10 June 2014, IEBR 3731 (Field No. CB 2015.11), adult female, collected by T.Q.
Nguyen on 22 April 2015, the same data as the holotype.
Species name 12S 16S Voucher number Reference
Babina daunchina KF185029 KF185065 HNNU20060103 Chen et al. 2013
Odorrana chapaensis DQ283372 DQ204475 AMNH A161439,
ROM 30943
Frost et al. 2006, Ngo et al.
2006
O. andersonii KF185021 KF185057 HNNU 001YN Chen et al. 2013
O. anlungensis KF185013 KF185049 HNNU 1008–109 Chen et al. 2013
O. daorum AF206101 AF206482 ROM 19053 Chen et al. 2005
O. grahami KF185015 KF185051 HNNU 1008–016 Chen et al. 2013
O. hmongorum EU861559 ROM 38605 Bain et al. 2009
O. jingdongensis KF185014 KF185050 20070711017 Chen et al. 2013
O. junlianensis KF185022 KF185058 HNNU 002 Chen et al. 2013
O. kuangwuensis KF184998 KF185034 HNNU 0908–185 Chen et al. 2013
O. lungshengensis KF185018 KF 185054 70028 Chen et al. 2013
O. margaretae KF184999 KF185035 HNNU 20050032 Chen et al. 2013
Odorrana mutschmanni sp. nov. KU356761 KU356765 IEBR 3723 This study
Odorrana mutschmanni sp. nov. KU356762 KU356768 IEBR 3725 This study
Odorrana mutschmanni sp. nov. KU356763 KU356766 IEBR 3727 This study
Odorrana mutschmanni sp. nov. KU356764 KU356767 IEBR 3730 This study
O. wuchuanensis KF185007 KF185043 HNNU 019 Chen et al. 2013
O. yizhangensis KF185012 KF185048 HNNU 1008–075 Chen et al. 2013
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TABLE 2. Uncorrected (“p”) distance matrix showing percentage pair wise genetic divergence (12S and 16S) between
members of the Odorrana andersonii species group.
Diagnosis. The new species was strongly supported as a member of Odorrana based on molecular analyses
(Fig. 1) and is distinguishable from its congeners by a combination of the following morphological characters: (1)
size large (SVL 85.9–91.6 mm in males, 108.7–110.1 mm in females); (2) head longer than wide; (3) vomerine
teeth present; (4) external vocal sacs absent; (5) snout short (SL/SVL 0.16–0.17); (6) tympanum large (TD/ED 0.70
in males, 0.68 in females); (7) dorsal surface of head and anterior part of body smooth, posterior part of body and
flanks with small tubercles; (8) supratympanic fold present; (9) dorsolateral fold absent; (10) webbing formula I0–
0II0–0III0–1/2IV1/2–0V; (11) in life, dorsum green with dark brown spots; (12) flanks greyish brown with dark
brown spots; (13) throat and chest grey, underside of limbs with large dark brown spots, edged in white, forming a
network.
Description of holotype. Adult male; SVL 85.9 mm; head longer than wide (HL 33.6 mm, HW 29.9 mm);
snout round anteriorly in dorsal view, projecting beyond lower jaw; nostril lateral, closer to the snout tip than to eye
(NS 6.2 mm, EN 7.5 mm); canthus rostralis distinct; pupil horizontally oval; loreal region slightly concave and
oblique; snout length greater than eye diameter (SL 14.0 mm, ED 9.9 mm); internarial distance wider than
interorbital distance and upper eyelid (IND 10.5. mm, IOD 9.7 mm, UEW 6.6 mm); tympanum distinct, round,
70% eye diameter (TD 6.9 mm); vomerine teeth in two oblique ridges; tongue cordiform, deeply notched
posteriorly; vocal sac absent.
Forelimbs: Forelimb length (FLL 16.6 mm), hand length (HAL 44.4 mm); relative finger lengths: II<I<IV<III;
finger webbing rudimental; tips of fingers expanded into discs, with circummarginal grooves, width of finger III
disc > 2 times of the width of phalanges and about 40% the diameter of tympanum; subarticular tubercles round,
formula 1, 1, 2, 2; inner metatarsal tubercle oval, elongate; outer metatarsal tubercle small; finger I with nuptial
pad, elongate.
Hindlimbs: Tibia longer than thigh (FeL 38.1 mm, TbL 46.9 mm), approximately five times longer than wide
(TbW 9.8 mm); tips of toes expanded into discs, with circummarginal grooves; width of toe IV disc narrower than
width of finger III disc, approximately two times of the width of phalanges; relative length of toes: I<II<III<V<IV;
webbing formula I0–0II0–0III0–1/2IV1/2–0V; subarticular tubercles prominent, formula 1, 1, 2, 3, 2; inner
metatarsal tubercle elongate; outer metatarsal tubercle absent.
Skin: Dorsal surface of head and anterior part of body smooth; posterior part of body and flanks with tubercles;
spinules present on lateral sides of body, anterior and posterior edge of tympanum; supratympanic fold present;
dorsolateral fold absent; dorsal surface of limbs granular; throat, chest, belly and ventral surface of thigh smooth.
Coloration in life: Iris black; dorsum green with dark brown spots; lateral side of head and flanks greyish
brown with dark brown spots; lips with dark bars; tympanum dark brown; spinules on flank ivory; dorsal surface of
fore and hindlimbs greyish brown with dark crossbars; throat and chest grey; ventral surface of fore and hindlimbs
and belly with large dark brown spots, edged in white, forming a network; toe webbing dark brown.
Species name 1 2 3 4567 8910111213
1. O. andersonii -
2. O. anlungensis 6.9 -
3. O. daorum 2.2 6.9 -
4. O. grahami 1.8 6.6 1.0 -
5. O. hmongorum 2.5 7.2 0.2 0.7 -
6. O. jingdongensis 2.0 6.7 1.9 1.7 2.2 -
7. O. junlianensis 1.7 6.8 1.2 0.3 1.0 1.8 -
8. O. kuangwuensis 3.0 6.8 3.4 2.8 3.3 2.8 2.8 -
9. O. lungshengensis 6.4 5.4 6.7 6.2 7.0 5.8 6.4 6.6 -
10. O. margaretae 2.9 7.1 3.4 2.7 3.9 2.5 2.9 2.8 6.4 -
11. Odorrana mutschmanni sp. nov. 5.8 7.4–7.5 5.7–5.8 5.8 6.4 5.6 5.9–6.0 6.1 7.2 6.0 -
12. O. wuchuanensis 4.8 7.4 4.2 4.5 4.4 4.6 4.7 5.4 6.6 5.6 4.3 -
13. O. yizhangensis 6.5 5.7 6.5 6.0 7.0 5.7 6.3 6.3 4.4 6.1 7.2 6.2 -
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Coloration in alcohol: Dorsum brown, flank cream with large spots; lips with black bars; fore and hindlimbs
brown with dark crossbars; throat and chest grey; belly and ventral surface of fore and hindlimbs with large black
spots; toe webbing brown.
Vari a t i o n. Measurements and morphological characters of the type series are given in Table 3.
FIGURE 1. The maximum parsimony tree based on partial 12S and 16S genes. This is one of six most parsimonious trees (TL
= 751; CI = 0.70; RI = 0.69). The dataset includes 1836 aligned characters of which 225 are parsimony informative. Numbers
above and under branches are MP/ML bootstrap values and Bayesian posterior probabilities (>50%), respectively. The arrows
show alternative placements supported by ML and BA analyses with the numbers above and below corresponding to the BP
and PP values, respectively. Asterisk denotes 100% value.
Sexual dimorphism. The males are smaller than the females in size (SVL: 85.9–91.6 mm, n = 8 vs 108.7–
110.1 mm, n = 3, respectively).
Etymology. We name this new species in honor of Dr. Frank Mutschmann, director of EXOMED in Berlin, in
recognition of his support of our research and conservation work in Vietnam. As common names we suggest
Mutschmann’s Frog (English), Ếch đá mut-x-man (Vietnamese), and Mutschmanns Frosch (German).
Ecological notes. Odorrana mutschmanni sp. nov. appears closely associated with karst environments.
Specimens were found at night between 19:00 and 23:00 h around a water pool near Coong Village, Duc Quang
Commune (Fig. 4). The surrounding habitat was a secondary karst forest of medium and small hardwoods mixed
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               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
  ……continued on the next page
Zootaxa 4084 (3) © 2016 Magnolia Press
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A NEW ODORRANA FROM VIETNAM
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 
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        
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               
          
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
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with shrubs and vines. Most of the specimens were found on rock boulders, ca. 0.5–1.0 m above the ground, few
frogs were in the water. Air temperature was 23.4–29.1
o
C and relative humidity was 57–79%. Other amphibian
species were found at the site, including Rhacophorus kio Ohler & Delorme, Polypedates mutus (Smith), and
Kurixalus bisacculus (Taylor).
FIGURE 2. Dorsolateral view (A) and ventral view (B) of the holotype (IEBR 3723, male) of Odorrana mutschmanni sp. nov.
in life.
Zootaxa 4084 (3) © 2016 Magnolia Press
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429
A NEW ODORRANA FROM VIETNAM
FIGURE 3. Ventral side of head and chest (A) and foot (B) of the holotype (IEBR 3723) of Odorrana mutschmanni sp. nov. in
preservative.
Distribution. Odorrana mutschmanni is currently known only from the type locality in Cao Bang Province,
northeastern Vietnam (Fig. 5). This species is presumably also to be found in adjacent karst formations in Ha Giang
Province of Vietnam and Guangxi and Yunnan provinces of China.
Comparisons. Within the Odorrana andersonii group (O. andersonii, O. jingdongensis, O. kwangwuensis, O.
margaretae, O. grahami, O. junlianensis, and O. wuchuanensis), Odorrana mutschmanni sp. nov. differs from O.
andersonii by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.5 in males and 0.45 in
females in O. andersonii), different ventral color pattern (large dark spots vs. immaculate white in O. andersonii),
males without spines on chest (vs. present in O. andersonii), the disc of finger III > 2 times base of phalanges (vs.
≤ 2 times base of phalanges in O. andersonii), and different egg color (wholly unpigmented vs. pigmented in O.
andersonii); from O. jingdongensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.54
in males and 0.51 in females in O. jingdongensis), different ventral color pattern (large dark spots vs. immaculate
white in O. jingdongensis), males without spines on chest (vs. present in O. jingdongensis), and the disc of finger
III > 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. jingdongensis); from O. margaretae by
having large dark spots on belly (vs. small dark spots in O. margaretae), males without spines on chest (vs. present
in O. margaretae), the disc of finger III > 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O.
margaretae), more developed toe webbing (complete to disc on I vs. as narrow fringe to disc on I in O.
margaretae), and different egg color (wholly unpigmented vs. pigmented in O. margaretae); from O. kuangwuensis
by having a larger body size (SVL 87–92 mm in males and 108–110 mm in females vs. 57 mm in males and 69–71
mm in females in O. kwangwuensis), a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.55 in males
and 0.5 in females in O. kwangwuensis), different ventral color pattern (large black spots vs. white with some black
spots in O. kwangwuensis), and the disc of finger III > 2 times base of phalanges (vs. ≤ 2 times base of phalanges in
O. kwangwuensis); from O. grahami by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.53
in males and 0.48 in females in O. grahami), different ventral color pattern (large black spots vs. immaculate white
in O. grahami), males without spines on chest (vs. present in O. grahami), the disc of finger III > 2x base of
phalanges (vs. finger III without disc in O. grahami), and different egg color (wholly unpigmented vs. pigmented
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in O. grahami); from O. junlianensis by having a higher ratio of TD/ED 0.70 in males and 0.68 in females (vs. 0.47
in males and 0.46 in females in O. junlianensis), the absence of external vocal sacs (vs. present in O. junlianensis),
and males without spines on chest (vs. white spinules present on the chest in O. junlianensis), the disc of finger III
> 2 times base of phalanges (vs. ≤ 2 times base of phalanges in O. junlianensis), and different egg color (wholly
unpigmented vs. pigmented in O. junlianensis); from O. wuchuanensis by having a larger size (SVL 87–92 mm in
males and 108–110 mm in females vs. 71–77 mm in males and 76–90 mm in females in O. wuchuaensis), a smaller
ratio TD/ED (0.70 in males and 0.68 in females vs. 0.83 in males and 0.8 in females in O. wuchuaensis), dorsal
surface of head and anterior part of body smooth (vs. shagreened in O. wuchuanensis), different color pattern of
flank and limbs (brown vs. green in O. wuchuanensis), and males without white spines on dorsal surface of arm
(vs. present in O. wuchuanensis).
Odorrana mutschmanni sp. nov. differs from O. absita, O. anlungensis, O. ammaiensis, O. aureola, O.
bacboensis, O. banaorum, O. bolavensis, O. chloronota, O. exiliversabilis, O. fengkaiensis, O. gigatympana, O.
graminea, O. hainanensis, O. heatwolei, O. hejingensis, O. hosii, O. huanggangensis, O. khalam, O. lipuensis, O.
lungshengensis, O. macrotympana, O. monjerai, O. morafkai, O. nanjingensis, O. narina, O. nasica, O. nasuta, O.
orba, O. rotodora, O. schmakeri, O. sinica, O. tiananensis, O. tianmuii, O. tormota, O. yentuensis, O.
yizhangensis, and O. zhaoi, by having a larger size (SVL 87–92 mm in males vs. ≤ 70 mm in males in other
species).
Odorrana mutschmanni sp. nov. further differs from O. absita, O. amamiensis, O. aureola, O. banaorum, O.
chloronota, O. exiliversabilis, O.gigatympana, O. graminea, O. hosii, O. indeprensa, O. khalam, O. leporipes, O.
livida, O. morafkai, O. nasica, O. nasuta, O. orba, O. swinhoana, O. tormota, O. trankieni, O. versabilis, O.
yentuensis, and O. zhaoi by the presence of black bars on lips (vs. absent in the latter).
Odorrana mutschmanni sp. nov. differs from O. absita, O. alungensis, O. banaorum, O. bolavensis, O.
exiliversabilis, O. hosii, O. indeprensa, O. khalam, O. leporpes, O.monjerai, O. narina, O. nasica, O. supranarina,
O. tormota, O. trankieni, O. utsunomiyaorum, O. versabilis, O. yentuensis, and O. zhaoi by lacking dorsolateral
folds (vs. present in the latter).
Odorrana muschmanni sp. nov. differs from O. amamiensis, O. bacboensis, O. bolavensis, O.chapaensis, O.
exiliversabilis, O. geminata, O. gigatypana, O. hainanensis, O. heatwolei, O. lipuensis, O. macrotympana, O.
nasica, O. orba, O. utsunomiyaorum, O. supranarina, O. tiannanensis, O. tormota, O. versabilis, and O. yentuensis
by having a green dorsum (vs. brown, light brown, olive-brown, reddish-brown, gray-blue, iridescent blue or grass-
green in the latter).
Odorrana mutschmanni sp. nov. differs from O. absita, O. aureola, O. bacboensis, O. banaorum, O.
bolavensis, O. chapaensis, O. chloronota, O. fengkaiensis, O. geminata, O. graminea, O. gigatympana, O.
heatwolei, O. indeprensa, O. ishikawae, O. khalam, O. lungshengensis, O. morafkai, O. nasica, O. orba, O.
swinhoana, O. tinananensis, O. tormota, O. trankieni, O. yentuensis, O. yizhangensis, and O. zhaoi by the absence
of vocal sacs in males (vs. present in the latter)
Discussion
Although a number of Odorrana species have been discovered in recent years, the diversity of the genus is not
fully understood. Two new species of the genus were just recently described in southern China, i.e., O. lipuensis
and O. fengkaiensis (Mo et al. 2015; Wang et al. 2015), and the discovery of this additional new species in Vietnam
suggests that the current species richness of the genus remains underestimated. Species complexes pose particular
problems to alpha taxonomy and species conservation. More studies using an integrative approach, i.e., combining
morphological and molecular data, will help to reveal the extent of species richness of Odorrana in the poorly
studied regions of northern Vietnam, in order to allow accurate inference of their phylogenetic relationships within
the morphologically complex species group.
The limestone karst forests of Cao Bang Province are being recognized as a center for new discoveries in
northeastern Vietnam. Besides O. mutschmanni, numerous new species of reptiles and amphibians have been
recently described from Ha Lang and Trung Khanh districts of Cao Bang Province, viz. Protobothrops
trungkhanhensis Orlov, Ryabov & Nguyen, 2009, Gracixalus waza Nguyen, Le, Pham, Nguyen, Bonkowski &
Ziegler, 2012, Oligodon nagao David, Nguyen, Nguyen, Jiang, Chen, Teynié & Ziegler, 2012, Gekko adleri
Nguyen, Wang, Yang, Lehmann, Le, Ziegler & Bonkowski, 2013, and Hemiphyllodactylus zugi Nguyen, Lehmann,
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Le, Duong, Bonkowski & Ziegler, 2013 (Orlov et al. 2009; David et al. 2012; Nguyen et al. 2012; Nguyen et al.
2013a,b). This study underpins that karst forests in Ha Lang District of Cao Bang Province still harbor
undiscovered new vertebrate species. However, biodiversity of karst forests in Cao Bang Province are currently
threatened due to quarrying for cement and road construction, expanding agriculture, and illegal timber logging.
Therefore, establishment of a new protected area (e. g., landscape and habitat protection area) should be considered
by provincial authorities to protect the remaining karst forests and its biodiversity in north-eastern Cao Bang
Province.
FIGURE 4. Habitat of Odorrana mutschmanni sp. nov. in Ha Lang District, Cao Bang Province, Vietnam.
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FIGURE 5. Map showing the type locality (black square) of Odorrana mutschmanni sp. nov. in Ha Lang District, Cao Bang
Province, Vietnam.
Acknowledgements
We are grateful to the directorates of the Forest Protection Department of Cao Bang Province and Ha Lang District
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for support of our field work and issuing relevant permits (Permit No. 216/UBND-NC). We thank H.T. An (Hanoi),
S. Herbst, T. Lehmann, M. Bernardes and M. van Schingen (Cologne) for their assistance in the field, and H.T. Ngo
for laboratory assistance. We thank E. Sterling (New York) and K. Koy (Berkeley) for providing the map. Thanks
to J.J. Rowley and two anonymous reviewers for their helpful comments. For the fruitful cooperation within joint
amphibian projects we cordially thank C.X. Le, T.H. Tran (IEBR, Hanoi) and T. Pagel and C. Landsberg
(Cologne). Field surveys in Cao Bang Province were partially funded by the Nagao Natural Environment
Foundation (Japan), the Deutsche Foschungsgemeinschaft (DFG)—International Collaboration Grant (BO 1907/
11–1), and the Cologne Zoo (Germany). This research was partially funded by the Institute of Ecology and
Biological Resources, VAST (IEBR.DT 03/15-16). Research of T. Q. Nguyen in Germany and equipment were
funded by the Alexander von Humboldt Foundation (VIE 1143441).
References
Bain, R.H. & Stuart, B.L. (2006 "2005") A new species of cascade frog (Amphibia: Ranidae) from Thailand, with new data on
Rana banaorum and Rana morafkai. Natural History Bulletin of the Siam Society, 53, 3–16.
Bain, R.H. & Stuart, B.L. (2006) Significant new records of the Junlian Odorous Frog, Odorrana junlianensis Huang, Fei, and
Ye, 2001. Hamadryad, 30, 151–156.
Bain, R.H., Lathrop, A., Murphy, R.W., Orlov, N.L. & Ho, T.C. (2003) Cryptic species of a cascade frog from Southeast Asia:
taxonomic revisions and descriptions of six new species. American Museum Novitates, 3417, 1–60.
Bain, R.H., Stuart, B.L., Nguyen, Q.T., Che, J. & Rao, D. (2009) A new Odorrana (Amphibia: Ranidae) from Vietnam and
China. Copeia, 2, 348–362.
Chen, L.Q., Murphy, R.W., Lathrop, A., Ngo, A., Orlov, N.L., Ho, C.T. & Somorjai, I.L.M. (2005) Taxonomic chaos in Asian
ranid frogs: An initial phylogenetic resolution. Herpetology Journal, 15, 231–243.
Chen, X., Chen, Z., Jiang, J., Qiao, L., Lu, Y., Zhou, K., Zheng, G., Zhai, X. & Liu, J. (2013) Molecular phylogeny and
diversification of genus Odorrana (Amphibia, Anura, Ranidae) inferred from two mitochondrial genes. Molecular
Phylogenetics and Evolution, 69, 1196-1202.
Chen, X., Zhou, K. & Zheng, G. (2010a) A new species of the genus Odorrana from China (Anura, Ranidae). Acta
Zootaxonomica Sinica, 35, 206–211. [In Chinese with English abstract]
Chen, X., Zhou, K. & Zheng, G. (2010b) A new species of Odor Frog from China (Anura: Ranidae). Journal of Beijing Normal
University (Natural Science), 46, 606–609. [In Chinese with English abstract]
David, P., Nguyen, T.Q., Nguyen, T.T., Jiang, K., Chen, T.-B., Teynié, A. & Ziegler, T. (2012) A new species of the genus
Oligodon Fitzinger, 1826 (Squamata: Colubridae) from northern Vietnam, southern China and central Laos. Zootaxa,
3498, 45−62.
Fei, L., Hu, S., Ye, C. & Huang, Y. (2009) Fauna Sinica. Amphibia. Volume 3. Anura. Science Press, Beijing, 887 pp
Fei, L., Ye, C. & Jiang, J. (2010) Colored Atlas of Chinese Amphibians. Sichuan Publishing House of Science & Technology,
Sichuan, 519 pp.
Frost, D.R. (2015) Amphibian Species of the World: an on-line reference. Version 6.0. American Museum of Natural History,
New York, USA. Available from: http://research.amnh.org/herpetology/amphibia/ (accessed 15 July 2015)
Frost, D.R., Grant, T., Faivovich, J., Bain, R., Haas, A., Haddad, C.F.B., Sa, R.O., Channing, A., Wilkinson, M., Donnellan,
S.C., Raxworthy, C., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. &
Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–291.
Giribet, G. & Wheeler, W. (1999) On gaps. Molecular Phylogenetics and Evolution, 13, 132–143.
Glaw, F. & Vences, M. (2007) A field guide to the amphibians and reptiles of Madagascar. Third Edition, Frosch Verlag,
Cologne.
Hedges, S.B. (1994) Molecular evidence for the origin of birds. Proceedings of the National Academy of Sciences of the United
States of America, 91, 2621–2624.
Kocher, T.D., Thomas, W.K, Meyer, A., Edwards, S.V., Pääbo, S., Villablanca, F.X. & Wilson, A.C. (1989) Dynamics of
mitochondrial DNA evolution in animals: Amplification and sequencing with conserved primers. Proceedings of the
National Academy of Sciences of the United States of America, 86, 6196−6200.
Le, M., Raxworthy, C.J., McCord, W.P. & Mertz, L. (2006) A molecular phylogeny of tortoises (Testudines: Testudinidae)
based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 40, 517−531.
http://dx.doi.org/10.1016/j.ympev.2006.03.003
Li, P., Lu, Y. & Rao, D. (2008) A new species of Cascade Frog (Amphibia, Ranidae) from Tibet, China. Acta Zootaxonomica
Sinica, 33, 537–541.
Liang, F., Chang, Y. & Cheng, L. (2001) Descriptions of two new species of the genus Odorrana in China. Acta
Zootaxonoomica Sinica, 26, 108–114.
Maeda, N. & Matsui, M. (1990) Frogs and Toads of Japan. Second Edition. Bun-Ichi Sogo Shuppan Co., Tokyo, Japan.
Matsui, M. (1994) A taxonomic study of the Rana narina complex, with description of three new species (Amphibia: Ranidae).
PHAM ET AL.
434
·
Zootaxa 4084 (3) © 2016 Magnolia Press
Zoological Journal of the Linnean Society, 111, 385–415.
Matsui, M. & Jaafar, I. (2006) A new cascade frog of the subgenus Odorrana from Peninsular Malaysia. Zoological Science,
23, 647–651.
Mo, Y., Chen, W., Wu, H., Zhang, W. & Zhou, S. (2015) A new species of Odorrana inhabiting complete sarkness in a karst
cave in Guangxi, China. Asian Herpetological Research, 6, 11–17.
Ngo, A., Murphy, R.W., Liu, W., Lathrop, A. & Orlov, N.L. (2006) The phylogenetic relationships of the Chinese and
Vietnamese Waterfall frogs of the genus Amolops. Amphibia-reptilia, 27, 81-92.
Nguyen, S.V., Ho, C.T. & Nguyen, T.Q. (2009) Herpetofauna of Viet Nam. Edition Chimaira, Frankfurt am Main.
Nguyen, T.Q., Le, M.D., Pham, C.T., Nguyen, T.T., Bonkowski, M. & Ziegler, T. (online: 2012/printed 2013) A new species of
Gracixalus (Amphibia, Anura, Rhacophoridae) from northern Vietnam. Organisms Diversity & Evolution, 13, 203–214.
Nguyen, T.Q., Lehmann, T., Le, M.D., Duong, T.H., Bonkowski, M. & Ziegler, T. (2013a) A new species of
Hemiphyllodactylus (Reptilia: Gekkonidae) from northern Vietnam. Zootaxa, 3736 (1), 89–98.
http://dx.doi.org/10.11646/zootaxa.3736.1.5
Nguyen, T.Q., Wang, Y-Y., Yang, Y-H, Lehmann, T., Le, M.D., Ziegler, T. & Bonkowski, M. (2013b) A new species of the
Gekko japonicus group (Squamata: Sauria: Gekkonidae) from the border region between China and Vietnam. Zootaxa,
3652 (5), 501–518.
http://dx.doi.org/10.11646/zootaxa.3652.5.1
Orlov, N.L., Ananjeva, N.B. & Ho, C. T. (2006) A new cascade frog (Amphibia: Ranidae) from central Vietnam. Russian
Journal of Herpetology, 13, 155–163.
Orlov, N.L., Le, N.N. & Ho, T.C. (2003) A new species of Cascade Frog from North Vietnam (Ranidae, Anura). Russian
Journal of Herpetology, 10, 123–134.
Orlov, N.L., Ryabov, S.A. & Nguyen, T.T. (2009) Two new species of genera Protobothrops Hoge et Romano-Hoge, 1983 and
Viridovipera Malhotra et Thorpe, 2004 (Ophidia: Viperidae: Crotalinae) from karst region in northeastern Vietnam. Part I.
Description of a new species of Protobothrops genus. Russian Journal of Herpetology, 16, 69–82.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817−818.
http://dx.doi.org/10.1093/bioinformatics/14.9.817
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. &
Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology, 61, 539−542.
Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of
mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the
Entomological Society of America, 87, 651–701.
Stuart, B.L. & Bain, R.H. (2005) Three new species of spinule-bearing frogs allied to Rana megatympanum Bain, Lathrop,
Murphy, Orlov & Ho, 2003, from Laos and Vietnam. Herpetologica, 61, 478–492.
Stuart, B.L. & Chan-ard, T. (2005) Two new Huia (Amphibia: Ranidae) from Laos and Thailand. Copeia, 2, 279–289.
Stuart, B.L., Chuaynkern, Y., Chan-ard, T. & Inger, R.F. (2006) Three species of frogs and a new tadpole from eastern Thailand.
Fieldiana. Zoology, 1543, 1–10.
Stuart, B.L., Orlov, N.L. & Chan-ard, T. (2005) A new cascade frog (Amphibia: Ranidae) from Laos and Vietnam. Raffles
Bulletin of Zoology, 53, 125–131.
Swofford, D.L. (2001) PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods), version 4. Sinauer Associates,
Sunderland, Massachusetts.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: Flexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876−4882.
http://dx.doi.org/10.1093/nar/25.24.4876
Tran, T.T., Orlov, N.L. & Nguyen, T.T. (2008) A new species of Cascade Frog of Odorrana Fei, Yi et Huang, 1990 genus
(Amphibia: Anura: Ranidae) from Bac Giang Province (Yen Tu Mountain Range, northeast Vietnam). Russian Journal of
Herpetology, 15, 212–224.
Wang, Y.Y., Lau, M.N., Yang, J.H., Chen, G.L., Liu, Z.Y., Pang, H. & Liu, Y. (2015) A new species of the genus Odorrana
(Amphibia: Ranidae) and the first record of Odorrana bacboensis from China. Zootaxa, 3999 (2), 235–254.
http://dx.doi.org/10.11646/zootaxa.3999.2.4
Yang, D. (2008) Amphibia. In: Yang, D., & Rao, D. (Eds.), Amphibia and Reptilia of Yunnan. Yunnan Publishing Group
Corporation, pp. 12–152.
APPENDIX. Specimens examined.
Odorrana andersonii (n=4): Vietnam: Son La Province: Copia: IEBR A.2015.84–87 (Field No. SL 2015.39, 49, 120, 165).
Odorrana chapaensis (n=4): Vietnam: Lao Cai Province: Sa Pa: IEBR A.2015.88, A.2015.89 (Field No. LC 2014.7, LC
2014.8); Vietnam: Lai Chau Province: Tam Duong District: IEBR 2337, 2361.
Odorrana bacboensis (n=5): Vietnam: Bac Giang Province: Tay Yen Tu: IEBR A.2013.73, IEBR 3654, ZFMK 92848;
Vietnam: Cao Bang Province: Ha Lang District: IEBR A.2015.82, 2015.83 (CB 2012.39, CB 2012.40).
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Odorrana geminate (n=3): Vietnam: Cao Bang Provinve: Phia Oac: VNMN 04718–20.
Odorrana graminea (n=8): Vietnam: Bac Giang Province: Tay Yen Tu: IEBR 3649, 3650, VNMN 1346, ZFMK 92849;
Vietnam: Cao Bang Province: Ha Lang District: IEBR A. 2015.78–81 (CB 2012.5, 2012.8, 2012.55, 2012.151).
Odorrana graham (n=3): Vietnam: Lai Chau Province: Tam Duong District: IEBR 2317, 2370, 2371.
Odorrana morafkai (n=3): Vietnam: Thanh Hoa Province: Xuan Lien: IEBR 2015.55–57 (XL 2012.169, 172, 173).
Odorrana lipuensis (n=5): Vietnam: Cao Bang Province: Ha Lang District: IEBR A.2015.63–67.
Odorrana nasica (n=7): Vietnam: Thanh Hoa Province: Xuan Lien: IEBR A. 2015.72–75 (XL 2011.59–74); Vietnam: Lai
Chau Province: Tam Duong District: IEBR 2320, 3140, 3141.
Odorrana tiannanensis (n=3): Vietnam: Thanh Hoa Province: Xuan Lien: IEBR A.2015.90; Vietnam: Thanh Hoa Province: Pu
Hu: IEBR A.2015.91; Vietnam: Cao Bang Province: Phia Oac NR: VNMN 04721–23.
Odorrana trankieni (n=9): Vietnam: Hoa Binh Province: Thuong Tien: IEBR A. 2015.68–71 (HB 2011.1–4); Vietnam: Son La
Province: Phu Yen District: Muong Do: VNMN 0431, 0432, 0461–63.
Odorrana yentuensis (n=7): Vietnam: Bac Giang Province: Tay Yen Tu: IEBR A.2013.76, A. 2015.76, A. 2015.77 (BG
2012.38, 2012.40), VNMN 1320, 1340, ZFMK 92851, 92852.
... Most of the species inhabit mountain streams at elevations of 200-2000 m and can also be found on rocks or branches near the riverbed (Fei et al. 2009a;Frost 2020). Seven species from the genus Odorrana have recently been described using both molecular and morphological analyses (Chen et al. 2010a, b;Kuramoto et al. 2011;Mo et al. 2015;Wang et al. 2015;Pham et al. 2016;Li et al. 2018a). The phylogeny and diversity of Odorrana and the systematic status of taxa within the genus have been debated by taxonomists (Frost et al. 2006;Che et al. 2007;Fei et al. 2009a;Kurabayashi et al. 2010;Chen et al. 2013;Li et al. 2015). ...
... We compared the morphological characters between the new species and its congeners, based on literature values from the following species: Odorrana absita , O. amamiensis (Matsui, 1994), O. andersonii (Boulenger, 1882), O. anlungensis (Hu et al., 1973), O. aureola (Stuart et al., 2006), O. bacboensis , O. banaorum , O. bolavensis , O. cangyuanensis (Yang, 2008), O. chapaensis (Bourret, 1937), O. chloronota (Günther, 1876;Che et al. 2020), O. exiliversabilis (Fei et al., 2001b), O. fengkaiensis (Wang et al., 2015), O. geminata (Bain et al., 2009), O. gigatympana (Orlov et al., 2006), O. grahami (Boulenger, 1917), O. graminea (Boulenger, 1900), O. hainanensis (Fei et al., 2001a), O. hejiangensis (Deng & Yu, 1992), O. hosii (Boulenger, 1891), O. huanggangensis (Chen et al., 2010a), O. indeprensa , O. ishikawae (Stejneger, 1901), O. jingdongensis (Fei et al., 2001a), O. junlianensis , O. khalam , O. kuangwuensis (Hu et al., 1966), O. kweichowensis (Li et al., 2018a), O. leporipes (Werner, 1930), O. lipuensis (Mo et al., 2015), O. livida (Blyth, 1856), O. lungshengensis (Liu & Hu, 1962), O. macrotympana (Yang, 2008), O. margaretae (Liu, 1950), O. mawphlangensis (Pillai & Chanda, 1977), O. monjerai (Matsui & Jaafar, 2006), O. morafkai , O. mutschmanni (Pham et al., 2016), O. nanjiangensis (Fei et al., 2007b), O. narina (Stejneger, 1901), O. nasica (Boulenger, 1903), O. nasuta (Fei et al., 2001b), O. orba , O. rotodora (Yang, 2008), O. schmackeri (Boettger, 1892), O. sinica , O. splendida (Kuramoto et al., 2011), O. supranarina (Matsui, 1994), O. swinhoana (Boulenger, 1903), O. tianmuii (Chen et al. 2010b), O. tiannanensis (Yang & Li, 1980), O. tormota (Wu, 1977), O. trankieni , O. utsunomiyaorum (Matsui, 1994), O. versabilis (Liu & Hu, 1962), O. wuchuanensis (Wu et al., 1983), O. yentuensis (Tran et al., 2008) and O. yizhangensis (Fei et al., 2007a). ...
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A new species, Odorrana sangzhiensis sp. nov. , is described, based on five specimens from Sangzhi County, Zhangjiajie City, Hunan Province, China. Molecular phylogenetic analyses, based on mitochondrial 12S rRNA and 16S rRNA gene sequences, strongly support the new species as a monophyletic group nested into the O. schmackeri species complex. The new species can be distinguished from its congeners by a combination of the following characters: (1) body size medium (SVL: 42.1–45.1 mm in males, 83.3–92.7 mm in females); (2) dorsolateral folds absent; (3) tympanum diameter 1.53 times as long as the width of the disc of finger III in females; 2.3 times in males; (4) dorsal skin green with dense granules and sparse irregular brown spots; males with several large warts on dorsum; (5) two metacarpal tubercles; (6) relative finger lengths: I ≤ II < IV < III; (7) tibiotarsal articulation beyond the tip of the snout; (8) ventral surface smooth in females; throat and chest having pale spinules in adult males; (9) dorsal limbs green or yellow green with brown transverse bands; and (10) paired external vocal sacs located at corners of the throat, finger I with light yellow nuptial pad in males. This discovery increases the number of Odorrana species to 59 and those known from China to 37.
... The odorous frogs of the genus Odorrana Fei, Ye & Huang 1990 comprise a species-rich group of ranids that range widely through East and Southeast Asia, including the southern and eastern Himalayas (AmphibiaWeb 2020; Frost et al. 2020). Odorrana species are typical inhabitants of high-gradient streams, often located in mountain environments, although several taxa including O. mutschmanni Pham, Nguyen, Le, Bonkowski & Ziegler 2016, O. wuchuanensis (Xu 1983 and O. lipuensis Mo, Chen, Wu, Zhang & Zhou 2015 have been found to inhabit karstic limestone caves. Recent phylogenetic studies confirmed that Odorrana is monophyletic and consists of at least seven clades that currently harbor 59 species (Frost et al. 2020;Chen et al. 2013). ...
... More than half of these occur in southern China, which constitutes a diversity hotspot for the genus from which new species are regularly described (e.g. O. lipuensis and O. kweichowensis Li, Xu, Lv, Jiang, Wei & Wang 2018, Mo et al. 2015Pham et al. 2016;Li et al. 2018). Additional Odorrana taxa are expected to be discovered as research efforts in remote regions of southern China increase. ...
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Article
The Chinese-Myanmar border area forms part of a long-acknowledged biodiversity hotspot. This region is characterised by dramatic topography and diverse landscapes, which support a high degree of biodiversity and endemism that remains largely understudied. Based on recent survey efforts we here describe a new frog species of the genus Odorrana from this area. Found in Dulongjiang village, Yunnan, China, the new species differs from a morphological perspective from its congeners through the combination of a grass-green dorsum with black posterior spots, absence of dorsolateral folds, a distinct pineal body, presence of supratympanic folds, absence of outer metacarpal, metatarsal and supernumerary tubercles, and a pair of external subgular vocal sacs. From a molecular perspective, pairwise genetic distances of 4.8% to 11.6% for DNA sequences of the mitochondrial gene for 16S ribosomal RNA differentiate the new species from other Odorrana. Our mitochondrial phylogeny suggests that the new species is a deeply divergent genetic lineage clustering with the clade including Odorrana wuchuanensis, and the Odorrana andersonii and Odorrana margaretae groups. The discovery of this new species emphasizes the need for further herpetological studies in the China-Myanmar border region.
... Saikia et al. (2017) Odorrana aureola Cascades and fast-flowing streams Paired subgular Odorrana bacboensis Cascades and fast-flowing streams Paired subgular Odorrana banaorum Cascades and fast-flowing streams Paired subgular Odorrana bolavensis Forest around fast-flowing streams Paired subgular Odorrana cangyuanensis Cascades and fast-flowing streams Present, shape not specified Odorrana chapaensis Cascades and fast-flowing streams Paired subgular Yang (1991), , Odorrana chloronota Cascades and fast-flowing streams Paired subgular Boulenger (1882), Odorrana exiliversabilis Cascades and fast-flowing streams Present, shape not specified Odorrana fengkaiensis Forest around fast-flowing streams ? Wang et al. (2015) Odorrana geminata Cascades and fast-flowing streams Paired subgular Odorrana gigatympana Cascades and fast-flowing streams Paired subgular Pham et al. (2016) Odorrana grahami Forest around fast-flowing streams Paired subgular , Odorrana graminea Cascades and fast-flowing streams Paired subgular , Odorrana hainanensis Forest around fast-flowing streams Present, shape not specified Odorrana hejiangensis Forest around fast-flowing streams Paired subgular , Odorrana hosii Cascades and fast-flowing streams Paired subgular , , , Odorrana huanggangensis Cascades and fast-flowing streams Present, shape not specified Odorrana indeprensa Forest around fast-flowing streams Paired subgular Odorrana ishikawae Cascades and fast-flowing streams Paired subgular Odorrana jingdongensis Cascades and fast-flowing streams Paired subgular , Lu et al. (2008) Odorrana junlianensis Forest around fast-flowing streams Paired subgular , Odorrana khalam Forest floor, near small streams Paired subgular Odorrana kuangwuensis Forest around fast-flowing streams ? Wu & Li (2004) Odorrana kweichowensis Forest around streams of variable size Paired subgular Li et al. (2018) Odorrana leporipes Cascades and fast-flowing streams ? ...
... Cascades and fast-flowing streams Paired subgular , Odorrana lungshengensis Forest around mountain streams ? Lau & Zhao (2004) Odorrana macrotympana Cascades and fast-flowing streams Present, shape not specified Odorrana margaretae Cascades and fast-flowing streams Absent Odorrana mawphlangensis Forest around fast-flowing streams No external pouches present Mahony (2008) Odorrana monjerai Cascades and fast-flowing streams Paired subgular Matsui & Jaafar (2006) Odorrana morafkai Forest floor, near small streams Paired subgular Odorrana mutschmanni Karst forest ponds Absent Pham et al. (2016) Odorrana nanjiangensis Cascades and fast-flowing streams Present, shape not specified Odorrana narina Forest around fast-flowing streams Paired subgular , Odorrana nasica Cascades and fast-flowing streams Paired subgular Yang (1991), Odorrana nasuta Cascades and fast-flowing streams Present, shape not specified Odorrana orba Forest around fast-flowing streams Paired subgular Odorrana schmackeri Cascades and fast-flowing streams Paired subgular , Odorrana sinica Cascades and fast-flowing streams Present, shape not specified Odorrana splendida Cascades and fast-flowing streams ? IUCN SSC Amphibian Specialist Group (2015a). ...
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Cascades and fast-flowing streams impose severe restrictions on acoustic communication, with loud broadband background noise hampering signal detection and recognition. In this context, diverse behavioural features, such as ultrasound production and visual displays, have arisen in the evolutionary history of torrent-dwelling amphibians. The importance of the vocal sac in multimodal communication is being increasingly recognized, and recently a new vocal sac visual display has been discovered: unilateral inflation of paired vocal sacs. In the diurnal stream-breeding Hylodidae from the Atlantic forest, where it was first described, this behaviour is likely to be enabled by a unique anatomical configuration of the vocal sacs. To assess whether other taxa share this exceptional structure, we surveyed torrent-dwelling species with paired vocal sacs across the anuran tree of life and examined the vocal sac anatomy of exemplar species across 18 families. We found striking anatomical convergence among hylodids and species of the distantly related basal ranid genera Staurois, Huia, Meristogenys and Amolops. Ancestral character state reconstruction identified three new synapomorphies for Ranidae. Furthermore, we surveyed the vocal sac configuration of other anuran species that perform visual displays and report observations on what appears to be unilateral inflation of paired vocal sacs, in Staurois guttatus-an extremely rare behaviour in anurans.
... Due to morphologically similarity in the genus, species that were formally thought to be widespread are now being recognized as complexes of species with much narrower distributions (Wang et al., 2015). In Vietnam, Pham et al. (2016a, b) recently discovered a new species (Odorrana mutschmanni Pham, Nguyen, Le, Bonkowski & Ziegler, 2016) and a new country record (O. lipuensis Mo, Chen, Wu, Zhang & Zhou, 2015). ...
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Odorrana fengkaiensis was recorded for the first time from Vietnam based on a new amphibian collection from Cao Bang, Bac Kan, Bac Giang, Quang Ninh provinces. Sequences of the specimens from Vietnam were identical to those of the paratypes of O. fengkaiensis from China. In addition, O. trankieni was newly recorded from Hoa Binh and Bac Giang provinces, Northern Vietnam, including the first description of female individuals. Genetic divergence between O. trankieni from Vietnam and its sister species, O. nasuta from China, ranged from 1.42 to 2.66%. The intra-specific genetic distance within O. trankieni ranged from 0.68 to 1.88% between the northeastern and northwestern populations.
... Recent studies in the border region between Vietnam and China have revealed a number of new amphibian species, viz. Gracixalus waza (Nguyen et al. 2013a), Odorrana mutschmanni (Pham et al. 2016), Vietnamophryne orlovi (Poyarkov et al. 2018) and Megophrys caobangensis (Nguyen et al. 2020) from Cao Bang Province; Leptobrachella nyx (Ohler et al. 2011), Tylototriton ziegleri (Nishikawa et al. 2013), Rhacophorus larissae (Ostroshabov et al. 2013) and Limnonectes nguyenorum (McLeod et al. 2015) from Ha Giang Province; and Oreolalax sterlingae (Nguyen et al. 2013b), Leptobrachella botsfordi (Rowley et al. 2013), Gracixalus sapaensis (Matsui et al. 2017), Megophrys fansipanensis (Tapley et al. 2018), and M. hoanglienensis (Tapley et al. 2018) from Lao Cai and Yen Bai provinces. ...
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... The evergreen forests in the border region between China and Vietnam harbor a high level of herpetofaunal diversity with a remarkable number of new discoveries from both countries in the last decade (Yang and Rao, 2008, Nguyen et al., 2009, 2013a, 2013b, 2013c, 2013dPham et al., 2016aPham et al., , 2016bTapley et al., 2017;Frost, 2019). In the last five years, seven new species of amphibians have been described in the border region from the Vietnam side, viz. ...
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We describe Odorrana liboensis sp. nov., a new species from the Maolan National Nature Reserve, Libo County, Guizhou Province, China. Phylogenetic analyses based on DNA sequences of the mitochondrial 12S rRNA, 16S rRNA, and ND2 genes supported the new species as an independent lineage, closely related to O. lipuensis. The uncorrected genetic distances between the 12S rRNA and 16S rRNA in the new species and its closest congener, O. lipuensis, were 6.06% and 5.19%, respectively. The new species is distinguished from its congeners by a combination of the following morphological characters: (1) having medium body size, with the snout-vent length (SVL) of adult females approximately 1.2 times as long as males at 56.9 ± 1.0 (55.8-58.2 mm, n = 9) in females and 48.7 ± 1.2 (47.1-49.9 mm, n = 5) in males; (2) head length greater than width in males and females; (3) tympanum distinctly visible, greater than one-half the diameter of the eye; (4) eyes big and prominent, width of upper eyelid (UEW) approximately 3/4 of eye diameter (ED); (5) dorsolateral folds absent; (6) two metacarpal tubercles; (7) relative finger lengths: II < I < IV < III; (8) subarticular tubercles on fingers prominent: 1, 1, 2, 2; (9) one metatarsal tubercle; (10) tibiotarsal articulation reaching to between the eye and nostril when the leg is stretched forward; (11) toes with entire webbing to disks; (12) subarticular tubercles on toes prominent: 1, 1, 2, 3, 2; (13) dorsal surfaces of limbs with distinct brownish-black bands; (14) smooth, grass-green dorsum with irregular brown mottling; (15) venter smooth, lacking black spots; and (16) lacking pectoral spinules, lacking vocal sacs, and light white nuptial pad present on finger I in males. The new species is currently only known from the type locality.
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