A new species of Odorrana (Amphibia: Anura: Ranidae) from Vietnam

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 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.

Full text

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
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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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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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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.

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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).
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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.