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We describe a new species of Cyrtodactylus on the basis of four specimens collected from the limestone karst forest of Phu Yen District, Son La Province, Vietnam. Cyrtodactylus sonlaensis sp. nov. is distinguished from the remaining Indochinese bent-toed geckos by a combination of the following characters: maximum SVL of 83.2 mm; dorsal tubercles in 13–15 irregular rows; ventral scales in 34–42 rows; ventrolateral folds prominent without interspersed tubercles; enlarged femoral scales 15–17 on each thigh; femoral pores 14–15 on each thigh in males, absent in females; precloacal pores 8, in a continuous row in males, absent in females; postcloacal tubercles 2 or 3; lamellae under toe IV 18–21; dorsal head with dark brown markings, in oval and arched shapes; nuchal loop discontinuous; dorsum with five brown bands between limb insertions, third and fourth bands discontinuous; subcaudal scales distinctly enlarged. In phylogenetic analyses, the new species is nested in a clade consisting of C. huongsonensis and C. soni from northern Vietnam and C. cf. pulchellus from Malaysia based on maximum likelihood and Bayesian analyses. In addition, we record Cyrtodactylus otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler for the first time from Son La Province based on specimens collected from Van Ho District.
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Accepted by A. Bauer: 29 Aug. 2017; published: 30 Oct. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4341 (1): 025
040
http://www.mapress.com/j/zt/
Article
25
https://doi.org/10.11646/zootaxa.4341.1.2
http://zoobank.org/urn:lsid:zoobank.org:pub:53DD68D9-1815-441B-B973-36A060F51475
A new species of Cyrtodactylus (Squamata: Gekkonidae) and the first record of
C. otai from Son La Province, Vietnam
TRUONG QUANG NGUYEN
1,2
, ANH VAN PHAM
3
, THOMAS ZIEGLER
4,5
,
HANH THI NGO
6
& MINH DUC LE
7,8,9,10
1
Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam.
E-mail: nqt2@yahoo.com
2
Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Cau Giay,
Hanoi, Vietnam
3
Faculty of Biology and Chemistry, Tay Bac University, Son La City, Son La Province, Vietnam. E-mail: phamanhdhsphn@gmail.com
4
AG Zoologischer Garten Köln, Riehler Strasse 173, D–50735 Cologne, Germany. E–mail: ziegler@koelnerzoo.de
5
Institute of Zoology, University of Cologne, Zülpicher Strasse 47b, D–50674 Cologne, Germany.
6
Faculty of Biology, Hanoi University of Science, Vietnam National University, 334 Nguyen Trai Road, Hanoi, Vietnam.
E-mail: ngothihanh.k56@hus.edu.vn
7
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
8
Central Institute for Natural Resources and Environmental Studies, Vietnam National University, 19 Le Thanh Tong, Hanoi, Vietnam
9
Department of Herpetology, American Museum of Natural History, Central Park West at 79
th
Street, New York, New York 10024
10
Corresponding author. E-mail: le.duc.minh@hus.edu.vn
Abstract
We describe a new species of Cyrtodactylus on the basis of four specimens collected from the limestone karst forest of
Phu Yen District, Son La Province, Vietnam. Cyrtodactylus sonlaensis sp. nov. is distinguished from the remaining Indo-
chinese bent-toed geckos by a combination of the following characters: maximum SVL of 83.2 mm; dorsal tubercles in
13–15 irregular rows; ventral scales in 34–42 rows; ventrolateral folds prominent without interspersed tubercles; enlarged
femoral scales 15–17 on each thigh; femoral pores 14–15 on each thigh in males, absent in females; precloacal pores 8, in
a continuous row in males, absent in females; postcloacal tubercles 2 or 3; lamellae under toe IV 18–21; dorsal head with
dark brown markings, in oval and arched shapes; nuchal loop discontinuous; dorsum with five brown bands between limb
insertions, third and fourth bands discontinuous; subcaudal scales distinctly enlarged. In phylogenetic analyses, the new
species is nested in a clade consisting of C. huongsonensis and C. soni from northern Vietnam and C. cf. pulchellus from
Malaysia based on maximum likelihood and Bayesian analyses. In addition, we record Cyrtodactylus otai Nguyen, Le,
Pham, Ngo, Hoang, Pham & Ziegler for the first time from Son La Province based on specimens collected from Van Ho
District.
Key words: Cyrtodactylus sonlaensis sp. nov., C. otai, molecular phylogeny, new record, taxonomy, Phu Yen, Van Ho.
Introduction
Son La Province is located in northwestern Vietnam and the province harbors a large area of 1,405,500 ha of
evergreen forest (The People's Committee of Son La Province 2007). However, the biodiversity of this province is
poorly studied, in particular reptiles and amphibians. Two new species were recently described from Son La
Province, namely Cyrtodactylus bichnganae Ngo & Grismer and Tylototriton anguliceps Le, Nguyen, Nishikawa,
Nguyen, Pham, Matsui, Bernardes & Nguyen (Ngo & Grismer 2010; Le et al. 2015a). In addition, ten new country
records of reptiles and amphibians have been reported from this province since 2010 (Le et al. 2014; 2015b,c;
Pham et al. 2014, 2016; Nguyen et al. 2015b).
During our recent field work in Son La Province, a series of bent-toed geckos was collected from Phu Yen and
NGUYEN ET AL.
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Moc Chau Districts. Morphological and molecular phylogenetic analyses revealed that the collection from Son La
Province contained two species: an unnamed species of Cyrtodactylus from Phu Yen District and a recently
described species, Cyrtodactylus otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler (Nguyen et al. 2015a),
from Van Ho District. We herein describe a new species of Cyrtodactylus and report the first record of C. otai from
Son La Province, Vietnam.
Material and methods
Sampling. Field surveys were conducted in Phu Yen and Van Ho districts, Son La Province, Vietnam, in June and
October 2016 (Fig. 1). Specimens were euthanized in a closed vessel with a piece of cotton wool containing ethyl
acetate (Simmons 2002), fixed in 85% ethanol and subsequently stored in 70% ethanol. Specimens were deposited
in the collections of the Institute of Ecology and Biological Resources (IEBR), Hanoi, Vietnam; the Faculty of
Biology and Chemistry, Tay Bac University (TBU), Son La Province, Vietnam; and the Vietnam National Museum
of Nature (VNMN), Hanoi, Vietnam.
Molecular data and phylogenetic analyses. We sequenced five new samples of Cyrtodactylus, four collected
from Son La Province and one from Nghe An Province (C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang
& Dau). We also included C. interdigitalis Ulber, C. cf. pulchellus Gray and available sequences from members of
the C. wayakonei species group (Fig. 2; clade B in Nguyen et al. 2015a). Since the sequences of C. pulchellus with
GenBank accession numbers HQ967203 and HQ967202 do not have precise localities, and recently described
species within the C. pulchellus species complex do not have the same gene region as that generated in this study
(Grismer et al. 2012), we were unable to assign the sequences to specific species within the complex. As a result,
we use C. cf. pulchellus for the sequences to reflect the uncertain nomenclature. Samples of C. elok Dring were
used as outgroups.
We used the protocols of Le et al. (2006) for DNA extraction, amplification, and sequencing. A fragment of the
mitochondrial gene, cytochrome c oxidase subunit 1 (COI), was amplified using the primer pair VF1-d and VR1-d
(Ivanova et al. 2006). After sequences were aligned by Clustal X v2 (Thompson et al. 1997), data were analyzed
using maximum parsimony (MP) and maximum likelihood (ML) as implemented in PAUP*4.0b10 (Swofford
2001) and Bayesian analysis (BA) as implemented in MrBayes v3.2 (Ronquist et al. 2012). 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 TIM+I+G for ML and combined Bayesian analyses
as selected by Modeltest v3.7 (Posada & Crandall 1998). The cutoff point for the burn-in function was set to 16 in
the Bayesian analysis, as -lnL scores reached stationarity after 16,000 generations in both runs. Nodal support was
evaluated using Bootstrap replication (BP) as estimated in PAUP and posterior probability (PP) in MrBayes v3.2.
Uncorrected pairwise divergences were calculated in PAUP*4.0b10 (Table 1).
Morphological characters. Measurements were taken with a digital caliper to the nearest 0.1 mm.
Abbreviations are as follows: snout-vent length (SVL), from tip of snout to anterior margin of cloaca; tail length
(TaL), from posterior margin of cloaca to tip of tail; trunk length or axilla-groin distance (AG), from posterior edge
of forelimb insertion to anterior edge of hindlimb insertion; head length (HL), from tip of snout to posterior margin
of the retroarticular; maximum head width (HW); maximum head height (HH), from occiput to underside of jaws;
greatest diameter of orbit (OD); snout to eye distance (SE), from tip of snout to anterior corner of orbit; eye to ear
distance (EyeEar), from anterior edge of ear opening to posterior corner of orbit; ear diameter (ED), maximum
diameter of ear; internarial distance (IND); maximum rostral width (RW); maximum rostral height (RH);
maximum mental width (MW); maximum mental length (ML); maximum body width at midbody (BW); forearm
length (ForeaL) from base of palm to elbow; crus length (CrusL), from base of heel to knee.
Scale counts were taken as follows: supralabials (SL); infralabials (IL); nasal scales surrounding nare (N, i.e.
nasorostral, supranasal, postnasals); postrostrals or internasals (IN); ciliaria (CIL), scales on eyelid fringe;
postmentals (PM); dorsal tubercle rows (DTR); granular scales surrounding dorsal tubercles (GST); ventral scales
in longitudinal rows at midbody (V); number of scales along the midbody from mental to anterior edge of cloaca
(SLB); enlarged femoral scales (EFS); femoral pores (FP); precloacal pores (PP); postcloacal tubercles (PAT);
number of subdigital lamellae on finger (NSF); number of subdigital lamellae on toe (NST). Bilateral scale counts
were given as left/right.
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A NEW CYRTODACTYLUS FROM VIETNAM
FIGURE 1. Map of sampling sites in northwestern Vietnam: 1) type locality of Cyrtodactylus sonlaensis sp. nov. in Phu Yen
District; 2) newly recorded locality of C. otai in Van Ho District, Son La Province; and 3) type locality of C. otai in Mai Chau
District, Hoa Binh Province.
FIGURE 2. Phylogram based on the Bayesian analysis. Number above and below branches are MP/ML bootstrap values and
Bayesian posterior probabilities (>50%), respectively. Asterisk represents 100% value.
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Results
Phylogenetic analyses. The final matrix consisted of 657 aligned characters, of which 217 were parsimony
informative. The alignment contained no gap. MP analysis of the dataset recovered a single most parsimonious tree
with 733 steps (CI = 0.52; RI = 0.73). In the ML analysis, the –Ln likelihood score of the single best tree found was
3939.12 after 2978 arrangements were tried. The topology derived from the Bayesian analysis (Fig. 2) was similar
to those in Nguyen et al. (2015a) and Le et al. (2016). C. chauquangensis was supported as a sister taxon to C.
vilaphongi Schneider, Nguyen, Le, Nophaseud, Bonkowski & Ziegler although this relationship received low
statistical values from all analyses. The major difference between the results based on different phylogenetic
analyses is the placement of C. cf. pulchellus Gray, which was recovered as a sister taxon to the new species with
strong support from both Bayesian and ML analyses. However, this placement was not corroborated by the MP
analysis, where the new species was grouped with C. bichnganae, C. huongsonensis, and C. soni in a clade with a
low support value (BP = 50%). In terms of genetic divergence, the new species was most similar to C.
huongsonensis Luu, Nguyen, Do & Ziegler and C. soni Le, Nguyen, Le & Ziegler with pairwise distance between
them ranging from 13.5–14.1% (Table 1). Sequences of two newly collected samples of C. otai from Son La
Province are identical to those of the type specimens from Hoa Binh Province (GenBank accession numbers
KT004370–71).
TABLE 1. Uncorrected (“p”) distance matrix showing percentage pairwise genetic divergence (COI) between new and
closely related species.
continued.
Species 1 2 3 4 5 6
C. bichnganae
C. bobrovi 16.3–16.4 –
C. cf. martini 15.8 16.9–17.1 –
C. chauquangensis 14.1 9.3 14.9 –
C. huongsonensis 14.6 15.8 16.0 14.6 –
C. otai 16.0 3.8 17.8 9.3 15.4 –
C. puhuensis 18.9 7.3–7.5 17.8 10.8 18.3 7.4
C. cf. pulchellus 20.3–21.0 18.6–19.2 18.9–20.1 18.1–18.8 17.1–18.1 19.2–19.6
C. sonlaensis sp. nov. 15.3–15.5 16.1–16.4 15.3–15.5 16.3–16.5 13.5–13.6 17.1–17.3
C. soni 13.4–13.9 16.7–16.9 16.4–16.7 14.7 5.0–.5.5 16.1–16.4
C. spelaeus 15.2–15.5 9.7–10.0 15.0–15.4 11.5–11.7 16.5–16.9 11.0–11.3
C. vilaphongi 15.7 9.4 15.5 8.3 15.2 9.4
C. wayakonei 14.8 16.6–16.8 6.7–6.9 14.0–15.0 16.6–16.9 18.1–18.4
Species 78910111213
C. bichnganae
C. bobrovi
C. cf. martini
C. chauquangensis
C. huongsonensis
C. otai
C. puhuensis
C. cf. pulchellus 18.9–19.7 –
C. sonlaensis sp. nov. 19.1–19.2 16.8–17.1
C. soni 19.1–19.5 18.5–18.6 13.9–14.1 –
C. spelaeus 11.3–11.7 18.6–19.0 15.3–15.7 15.4–16.1 –
C. vilaphongi 10.4 19.9–20.2 17.0–17.1 15.7–16.1 11.7–12.0 –
C. wayakonei 18.2–18.4 18.3–19.6 15.6–15.9 17.2–17.7 16.1–16.5 15.8–16.2 –
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A NEW CYRTODACTYLUS FROM VIETNAM
Cyrtodactylus sonlaensis sp. nov.
(Figs. 3–5)
Holotype. IEBR A.2017.1 (Field No. SL2016.68), adult male, collected on 18 June 2016 by A.V. Pham and H.V.
Tu, in the karst forest near Bang Village (21
o
05.700’N, 104
o
48.179’E, elevation 1050 m above sea level, asl.),
Muong Bang Commune, Phu Yen District, Son La Province, northwestern Vietnam.
Paratypes. TBU 2017.1 (Field No. SL2016.67, subadult male), IEBR A.2017.2 (Field No. SL2016.69, adult
female), the same data as the holotype; VNMN 2017.1 (Field No. SL2016.467, adult female), collected on 28
October 2016 by A.V. Pham and T.Q.L. Hoang, in the karst forest near Bang Village (21
o
06.406’N, 104
o
47.810’E,
elevation 890 m a.s.l.), Muong Bang Commune, Phu Yen District, Son La Province, northwestern Vietnam.
Diagnosis. The new species can be distinguished from other members of the genus Cyrtodactylus by a
combination of the following characters: medium size (SVL up to 83.2 mm); dorsal tubercles in 13–15 irregular
rows; 34–42 ventral scale rows; ventrolateral folds present without interspersed tubercles; 15–17 enlarged femoral
scales on each thigh; femoral pores 14 or 15 on each thigh of males, absent in females; precloacal pores 8, in a
continuous row in males, absent in females; postcloacal tubercles 2 or 3; lamellae under toe IV 18–21; dorsal head
with dark brown marking, oval and arched shape; nuchal loop discontinuous; five brown dorsal bands between
limb insertions, third and fourth discontinuous; subcaudal scales transversely enlarged.
FIGURE 3. The male holotype (IEBR A.2017.1) of Cyrtodactylus sonlaensis sp. nov. in life. Photo A. V. Pham.
FIGURE 4. Cloacal region of the holotype of Cyrtodactylus sonlaensis sp. nov. (IEBR A.2017.1). Photo A. V. Pham.
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FIGURE 5. The female paratype (VNMN 2017.1) of Cyrtodactylus sonlaensis sp. nov. in life. Photo A. V. Pham.
Description of holotype. Adult male, snout-vent length (SVL) 77.5 mm; body elongate (AG/SVL 0.46); head
distinguished from neck, elongate, depressed (HL/SVL 0.28, HW/HL 0.69, HH/HL 0.44); supranasals in contact
with each other anteriorly, separated from each other by a small scale posteriorly; nares oval, surrounded by
supranasal, rostral, first supralabial, and three postnasals; loreal region concave; snout long (SE/HL 0.41), round
anteriorly, longer than diameter of orbit (OD/SE 0.67); snout scales small, round, granular, larger than those in
frontal and parietal regions; eye large (OD/HL 0.28), pupils vertical; upper eyelid fringe with spinous scales; ear
opening oval, obliquely directed, small in size (ED/HL 0.09); rostral wider than high (RH/RW 0.71) with a medial
suture, bordered by first supralabial, nostril and supranasal on each side; mental triangular, as wide as rostral (RW
3.1 mm, MW 3.0 mm), wider than high (ML/MW 0.76); postmentals two, enlarged, in contact posteriorly, bordered
by mental anteriorly, first infralabial laterally, and an enlarged chin scale posteriorly; supralabials 11/11;
infralabials 11/9.
Dorsal scales granular; dorsal tubercles round, 3 or 4 times larger than the size of adjoining scales, conical,
present on occiput, back and tail base, each surrounded by 9–11 granular scales, in 14 or 15 irregular longitudinal
rows at midbody; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, round, subimbricate,
largest posteriorly, in 34–36 longitudinal rows at midbody; lateral skin folds distinct, without tubercles; gular
region with homogeneous smooth scales; ventral scales between mental and cloacal slit 189–193 (counted three
times); precloacal groove absent; three rows of enlarged scales present in posterior region of pore-bearing scales;
femoral pores bearing scales enlarged, in a continuous row with pore-bearing precloacal scales on the left side but
separated from pore-bearing precloacal scales by 2 poreless femoral scales on the right side; femoral pores 15/14;
precloacal pores eight, in a continuous row.
Fore and hind limbs moderately slender (ForeaL/SVL 0.17, CrusL/SVL 0.24); dorsal surface of forelimbs
covered by few slightly developed tubercles; dorsal surface of hind limbs covered by distinctly developed
tubercles; interdigital webbing weakly developed; subdigital lamellae: finger I 14/15 (with 6/6 basally broadened
lamellae), finger II 17/17 (7/7), finger III 16/18 (7/7), finger IV 18/17 (7/7), finger V 18/17 (8/7), toe I 15/16 (6/6),
toe II 18/17 (7/7), toe III 17/18 (7/7), toe IV -/18 (8/7), toe V 20/19 (8/7).
Tail complete, longer than snout-vent length (TaL 96.5 mm, Tal/SVL 1.24); postcloacal tubercles 3/3; dorsal
tail base with distinct tubercles; subcaudals distinctly enlarged, smooth.
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A NEW CYRTODACTYLUS FROM VIETNAM
TABLE 2. Morphological characters of Cyrtodactylus sonlaensis sp. nov. and C. otai from Son La Province, Vietnam
(measurements in mm, * = regenerated or broken tail, Min = minimum, Max = maximum, other abbreviations defined in
the text).
Cyrtodactylus sonlaensis sp.nov. Cyrtodactylus otai
IEBR A.2017.1
Holotype
TBU 2017.1
Paratype
IEBR A.2017.2
Paratype
VNMN 2017.1
Paratype
IEBR A.2017.3 TBU 2017.2
Sex Male Subadult male Female Female Male Female
SVL 77.5 63.1 71.2 83.2 92.5 88.5
TaL 96.5 33.3* 89.8 103.0 61* 60*
HL 22 18.4 20.5 22.6 24.0 23.7
HW 15.2 12.9 14.5 15.2 17.0 16.5
HH 9.8 8.2 9.5 9.6 11.2 10.1
OD 6.2 5.3 6.0 6.5 6.5 6.1
SE 9.2 8.5 8.7 9.7 10.0 10.1
EE 7.4 5.8 6.4 6.4 7.1 6.6
NE 6.8 5.3 6.3 7.0 7.0 7.1
ED 2.0 1.5 1.6 2.0 1.6 1.4
ForeaL 12.9 11.0 12.5 13.5 14.9 14.8
CrusL 18.4 13.0 15.5 17.9 17.6 17.6
AG 36 27.0 30.4 33.2 38.8 41.4
BW 13.4 11.0 12.8 13.5 15.0 15.4
IND 2.8 2.3 2.5 3.0 3.1 3.1
IOD 4.0 3.0 3.4 4.0 3.4 3.8
RW 3.1 3.0 3.2 3.5 3.6 3.7
RH 2.2 2.0 2.1 2.4 2.6 2.5
MW 3.0 3.0 3.1 3.4 3.5 3.5
ML 2.3 2.1 2.0 2.4 2.5 2.5
SPL 11/11 11/11 11/11 11/9 10/10 10/10
IFL 11/9 11/10 10/9 10/10 9/10 10/10
N 5/5 5/5 5/5 5/5 5/5 4/4
IN 0 2 0 0 0 0
CIL 26–29 26–28 27–28 26–28 28/29 26/25
PM 2 2 2 2 2 2
GST 9–11 9–10 9–10 9–10 9–10 8–9
V 34–36 40–42 37–39 34–37 41 40
SLB 189–193 200–202 196–201 195–200 185 186
FP 15/14 13/14 (pitted
scales)
000 0
PP 8 8 0 0 7 0
PAT 3/3 2/2 2/3 2/2 3/3 3/2
TubR 14–15 15 13 14–15 11–12 12–13
EFS 17/15 16/17 17/17 17/16 0 0
NSF
I 6+ 8/6+9 7+9/6+9 8+10/8+8 7+9/7+9 4+10/5+11 4+9/4+9
......continued on the next page
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Coloration in preservative. Ground color of dorsal head and back greyish brown; snout region yellowish with
three dark brown spots; dorsal head with dark brown marking, oval and arched shape; a dark stripe extending from
posterior corner of eye rearwards to above tympanum; labials brown with cream sutures; neck with some large dark
blotches, forming a discontinuous band anteriorly and a continuous band posteriorly; dorsum with five transverse
dark brown bands between fore and hind limb insertions, edged in white anteriorly and posteriorly, third and fourth
bands broken; dorsal surface of fore and hind limbs with dark brown blotches; tail greyish cream with ten dark
brown bands; chin, throat, chest, belly and lower limbs cream; ventral surface of tail grey with seven dark brown
bands. For coloration in life see Fig. 3.
Sexual dimorphism and variation. The females differ from male specimens in the absence of precloacal-
femoral pores and hemipenial swellings at the tail base. For other morphological characters see Table 2.
Distribution. Cyrtodactylus sonlaensis sp. nov. is currently known only from the type locality in Phu Yen
District, Son La Province, Vietnam (Fig. 1).
Etymology. The specific epithet “sonlaensis” refers to the type locality, Son La Province, where the new
species was discovered.
Natural history. Specimens were found at night between 19:00 and 21:00, on trees near limestone cliffs and in
rock crevices, about 0.1–2.0 m above the ground, at elevations between 900 and 1200 m a.s.l. The surrounding
habitat was disturbed evergreen karst forest of medium hardwood and shrub. The humidity was approximately 80–
90% and the air temperature ranged from 22 to 29
o
C.
Comparisons. We compared the new species with its congeners from Vietnam and neighboring countries in
mainland Indochina, including Laos, Cambodia, Myanmar and Thailand based on examination of specimens (see
Appendix) and data obtained from the literature (Smith 1917, 1921a,b, 1935; Taylor 1963; Ulber 1993; Bauer
2002, 2003; Bauer et al. 2002, 2003, 2009, 2010; Ziegler et al. 2002, 2010, 2013; Pauwels & Sumontha 2014;
Pauwels et al. 2004, 2013, 2014a,b, 2016; Nguyen et al. 2006, 2014; Hoang et al. 2007; Orlov et al. 2007; Grismer
& Ahmad; Ngo 2008; Ngo & Bauer 2008; Ngo & Grismer 2010, 2012; Ngo & Pauwels 2010; Ngo & Chan 2010,
2011; Ngo et al. 2008, 2010; Sumontha et al. 2010, 2012, 2014; Chan-ard & Makchai, 2011; David et al. 2011;
Schneider et al. 2011; Luu et al. 2011, 2014, 2015, 2016a,b,c; Grismer et al. 2012; Kunya et al. 2014, 2015;
Nazarov et al. 2014; Panitvong et al. 2014; Le et al. 2016; and Connette et al. 2017).
Cyrtodactylus sonlaensis sp. nov. has distinctly enlarged subcaudals, which are only slightly or not enlarged in
the following species: C. ayeyarwadyensis Bauer, C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen,
Hoang & Ziegler, C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. brevidactylus Bauer, C.
brevipalmatus (Smith), C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C.
buchardi David, Teynié & Ohler, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, C.
cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. dati
Ngo, C. gunungsenyumensis Grismer, Wood, Anuar, Davis, Cobos & Murdoch, C. gansi Bauer, C. hitchi Riyanto,
Kurniati & Engilis, C. huynhi Ngo & Bauer, C. irregularis (Smith), C. mandalayensis Mahony, C. martini Ngo, C.
TABLE 2. (Continued)
Cyrtodactylus sonlaensis sp.nov. Cyrtodactylus otai
IEBR A.2017.1
Holotype
TBU 2017.1
Paratype
IEBR A.2017.2
Paratype
VNMN 2017.1
Paratype
IEBR A.2017.3 TBU 2017.2
II 7 + 10/7+10 7+10/7+10 7+12/7+10 7+10/7+10 5+11/5+11 6+9/6+9
III 7 + 9/7+11 8+11/7+? 7+11/7+12 7+12/7+12 6+12/6+13 6+11/6+11
IV 7 + 11/7+10 8+11/8+10 7+10/7+12 7+12/7+12 7+12/7+12 6+12/6+12
V 8 + 10/7+10 7+9/7+10 7+8/7+11 6+11/6+10 5+12/5+11 5+9/5+11
NST
I 6+9/6+10 6+10/6+9 7+9/7+8 7+9/6+9 5+9/4+10 4+10/5+9
II 7+11 /7+10 8+10/8+9 7+10/9+10 8+9/8+10 6+11/6+11 5+10/6+10
III 7+10/7+11 7+13/9+11 7+11/7+12 9+11/8+12 5+14/6+14 5+14/6+14
IV 8+-/7+11 8+13/8+11 9+12/9+11 8+13/8+11 7+13/7+14 7+12/7+12
V 8+12/7+12 8+12/8+12 8+12/8+12 8+12/7+12 7+13/6+15 6+13/6+14
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A NEW CYRTODACTYLUS FROM VIETNAM
otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. papilionoides Ulber & Grossmann, C. phuocbinhensis
Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C.
pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, C. quadrivirgatus Tay lor, C. ranongensis Sumontha,
Pauwels, Panitvong, Kunya & Grismer, C. slowinskii Bauer, C. sommerladi Luu, Bonkowski, Nguyen, Le,
Schneider, Ngo & Ziegler, C. tamaiensis (Smith), C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop,
Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy & Zhang, C. thuongae Phung, Van Schingen, Ziegler
& Nguyen, C. vilaphongi Schneider, Nguyen, Duc Le, Nophaseud, Bonkowski & Ziegler, C. wakeorum Bauer, C.
wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, C. ziegleri Nazarov, Orlov, Nguyen & Ho.
Cyrtodactylus sonlaensis sp. nov. differs from C. aequalis Bauer by the absence of precloacal pores in females
(vs. 9), fewer dorsal tubercle rows (13–15 vs. 24), and more ventral scale rows (34–42 vs. 24); from C.
annandalei Bauer by its larger size (SVL 71.2–83.2 mm vs. 49.0–55.0 mm), more precloacal-femoral pores in
males (37 vs. 12–24), and fewer dorsal tubercle rows (13–15 vs. 16–18); from C. angularis (Smith) by having
more precloacal pores in males (8 vs. 3), the absence of precloacal pores in females (vs. 3), and the presence of
femoral pores in males (vs. absence); from C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad,
Bauer, Wangkulangkul, Grismer & Pauwels by having fewer dark caudal bands (10 vs. 13–14); from C.
auribalteatus Sumontha, Panitvong & Deein by having more enlarged femoral scales on each thigh (15–17 vs. 5–
7), more femoral pores on each thigh in males (14–15 vs. 4–5), and more precloacal pores in males (8 vs. 6); from
C. badenensis Nguyen, Orlov & Darevsky by having more ventral scale rows (34–42 vs. 25–29), the presence of
enlarged femoral scales (vs. absence), and the presence of precloacal-femoral pores in males (vs. absence); from C.
bichnganae Ngo & Grismer by its smaller size (SVL 71.2–83.2 mm vs. 95.3–99.9 mm), having more enlarged
femoral scales on each thigh (15–17 vs. 11–13), more femoral pores on each thigh in males (14–15 vs. 9), and
fewer precloacal pores in males (8 vs. 10); from C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler by having
more precloacal-femoral pores in males (37 vs. 34), more infralabials (9–11 vs. 8), and more ventral scales at
midbody (189–202 vs. 158–170); from C. calamei Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler by
having fewer postcloacal tubercles (2–3 vs. 4) and the absence of precloacal-femoral pores in females (vs. 38);
from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen by its smaller size (SVL 71.2–83.2 mm vs.
90.4–94.0 mm), having more enlarged femoral scales on each thigh (15–17 vs. 8), more femoral pores on each
thigh in males (14–15 vs. 6), fewer lamellae under finger IV (17–19 vs. 22) and under toe IV (18–21 vs. 23–25);
from C. chanhomeae Bauer, Sumontha & Pauwels by having more precloacal-femoral pores in males (37 vs. 32)
and the absence of precloacal-femoral pores in females (vs. 34); from C. chauquangensis Hoang, Orlov, Ananjeva,
Johns, Hoang & Dau by its smaller size (SVL 71.2–83.2 mm vs. 90.9–99.3 mm), the presence of enlarged femoral
scales (vs. absence), the presence of femoral pores in males (vs. absence), having more precloacal pores in males (8
vs. 6), and the absence of precloacal pores in females (vs. 7); from C. chrysopylos Bauer by having fewer
precloacal pores in males (8 vs. 10) and the presence of femoral pores in males (vs. absence); from C. condorensis
(Smith) by having more precloacal pores in males (8 vs. 4–7) and a different dorsal color pattern (banded vs.
blotched); from C. consobrinoides (Annandale) by its larger size (SVL 71.2–83.2 mm vs. 48.0 mm), more
precloacal pores in males (8 vs. 4), and more ventral scale rows (34–42 vs. 24–30); from C. cucphuongensis Ngo &
Chan, by its smaller size (SVL 71.2–83.2 mm vs. 96.0 mm), the presence of precloacal-femoral pores in males (vs.
absence), having fewer lamellae under finger IV (17–19 vs. 21), and under toe IV (18–21 vs. 24); from C. darevskii
Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov by having fewer precloacal-
femoral pores in males (37 vs. 38–44) and the absence of precloacal-femoral pores in females (vs. 24–34); from C.
doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri by its smaller size (SVL
reaching 83.2 mm vs. 90.5 mm), having more precloacal pores in males (8 vs. 6) and fewer dorsal tubercle rows
(13–15 vs. 19–20); from C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya by
having more precloacal-femoral pores in males (37 vs. 17–19), the absence of precloacal-femoral pores in females
(vs. 0–7), and more lamellae under finger IV (17–19 vs. 16); from C. eisenmanae Ngo by having fewer ventral
scale rows (34–42 vs. 44–45), more enlarged femoral scales on each thigh (15–17 vs. 4–6), and the presence of
precloacal-femoral pores in males (vs. absence); from C. erythrops Bauer, Kunya, Sumontha, Niyomwan,
Panitvong, Pauwels, Chanhome & Kunya by having more ventral scale rows (34–42 vs. 28), fewer precloacal pores
in males (8 vs. 9), more lamellae under finger IV (17–19 vs. 16), and a different dorsal color pattern (banded vs.
blotched); from C. feae Boulenger by its larger size (SVL 71.2–83.2 mm vs. 45.0 mm) and the presence of
precloacal-femoral pores in males (vs. absence); from C. grismeri Ngo by the presence of precloacal-femoral pores
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in males (vs. absence) and the presence of enlarged femoral scales (vs. absence); from C. hinnamnoensis Luu,
Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler by its smaller size (SVL reaching 83.2 mm vs. 100.6 mm) and
having fewer postcloacal tubercles (2–3 vs. 4–5); from C. hontreensis Ngo, Grismer & Grismer by having more
enlarged femoral scales on each thigh (15–17 vs. 2–5); from C. huongsonensis Luu, Nguyen, Do & Ziegler by its
smaller size (SVL reaching 83.2 mm vs. 89.8 mm), having more enlarged femoral scales on each thigh (15–17 vs.
7–9), more precloacal-femoral pores in males (37 vs. 23), and the absence of precloacal-femoral pores in females
(vs. 23); from C. interdigitalis Ulber by having fewer femoral pores on each thigh in males (14–15 vs. 16–18) and
fewer precloacal pores in males (8 vs. 14); from C. intermedius (Smith) by having more enlarged femoral scales on
each thigh (15–17 vs. 6–10), the presence of femoral pores (vs. absent), having fewer lamellae under finger IV (17–
19 vs. 20) and under toe IV (18–21 vs. 22); from C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu,
Sirisamphan & Pauwels by having fewer dorsal tubercle rows (13–15 vs. 18–20), more femoral pores on each thigh
in males (14–15 vs. 6), and more precloacal pores in males (8 vs. 5); from C. jaegeri Luu, Calame, Bonkowski,
Nguyen & Ziegler by having more ventral scale rows (34–42 vs. 31–32), fewer precloacal-femoral pores in males
(37 vs. 44), and the absence of precloacal-femoral pores in females (vs. 24); from C. jarujini Ulber, by its smaller
size (SVL 71.2–83.2 mm vs. 85.0–90.0 mm), having fewer precloacal-femoral pores in males (37 vs. 52–54),
generally more lamellae under finger IV (17–19 vs. 15–17), and a different dorsal color pattern (banded vs.
blotched); from C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov &
Chulisov by having fewer precloacal-femoral pores in males (37 vs. 40–44); from C. khelangensis Pauwels,
Sumontha, Panitvong & Varaguttanonda by having more precloacal pores in males (8 vs. 2–5), fewer dorsal
tubercle rows (13–15 vs. 16–20), more femoral pores on each thigh in males (14–15 vs. 6) and the absence of
precloacal pores in females (vs. 6); from C. kingsadai Ziegler, Phung, Le & Nguyen by having more enlarged
femoral scales on each thigh (15–17 vs. 9–12), more femoral pores on each thigh in the males (14–15 vs. 1–4), and
the absence of precloacal pores in females (vs. 4–8); from C. kunyai Pauwels, Sumontha, Keeratikiat &
Phanamphon by having more precloacal pores in males (8 vs. 3) and fewer dorsal tubercle rows (13–15 vs. 16–20);
from C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer &
Pauwels by the absence of precloacal-femoral pores in females (vs. 33–43); from C. lenya Mulcahy, Thura & Zug
by the presence of precloacal-femoral pores in males (vs. absence) and more ventral scale rows (34–42 vs. 29);
from C. lomyenensis Ngo & Pauwels by having fewer precloacal-femoral pores in males (37 vs. 39–40) and the
absence of precloacal-femoral pores in females (vs. 32); from C. macrotuberculatus Grismer & Ahmad by its
smaller size (SVL reaching 83.2 mm vs. 120.0 mm), more ventral scale rows (34–42 vs. 17–28) and fewer dorsal
tubercle rows (13–15 vs. 19–27); from C. multiporus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov,
Konstantinov & Chulisov by having fewer precloacal-femoral pores in males (37 vs. 58–60); from C. nigriocularis
Nguyen, Orlov & Darevsky by having more precloacal pores in males (8 vs. 0–2), the presence of enlarged femoral
scales (vs. absence), the presence of femoral pores in males (vs. absence), and a different dorsal color pattern
(banded vs. uniformly brown); from C. oldhami (Theobald) by having more precloacal pores in males (8 vs. 1–4),
the presence of femoral pores in males (vs. absence), and a different dorsal color pattern (banded vs. striped and
spotted); from C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler by having more precloacal pores
in males (8 vs. 4), the presence of enlarged femoral scales (vs. absence), the presence of femoral pores in males (vs.
absence), and the absence of precloacal pores in females (vs. 4); from C. payarhtanensis Mulcahy, Thura & Zug by
the presence of precloacal-femoral pores in males (vs. absence) and having more ventral scale rows (34–42 vs. 26–
32); from C. peguensis Boulenger by the presence of femoral pores in males (vs. absence) and a different dorsal
color pattern (banded vs. dark brown spots); from C. phetchaburiensis Pauwels, Sumontha & Bauer by its larger
size (SVL 71.2–83.2 mm vs. 57.5 mm), having more ventral scale rows (34–42 vs. 33), the presence of femoral
pores in males (vs. absence), more precloacal pores in males (8 vs. 5) and a different dorsal color pattern (banded
vs. blotched); from C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu by having fewer postcloacal
tubercles (2–3 vs. 4–5), more scale rows from mental to the front of cloacal slit (189–202 vs. 161–177); from C.
phuketensis Sumontha, Pauwels, Kunya, Nitikul, Samphanthamit & Grismer by its smaller size (SVL reaching 83.2
mm vs. 114.7 mm) and more ventral scale rows (34–42 vs. 22–24); from C. puhuensis Nguyen, Yang, Le, Nguyen,
Orlov, Hoang, Nguyen, Jin, Rao, Hoang, Che, Murphy & Zhang by having more precloacal pores in males (8 vs.
5), the presence of femoral pores in males (vs. absence), and fewer lamellae under toe IV (18–21 vs. 23); from C.
pulchellus Gray by its smaller size (SVL reaching 83.2 mm vs. 114.1 mm), more ventral scale rows (34–42 vs. 29–
34), and fewer dorsal tubercle rows (13–15 vs. 22–26); from C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu,
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A NEW CYRTODACTYLUS FROM VIETNAM
Dang, Dinh & Schmitz by having more enlarged femoral scales on each thigh (15–17 vs. 7–10), more precloacal-
femoral pores in males (37 vs. 20–28), and the absence of precloacal-femoral pores in females (vs. 17–22); from C.
rufford Luu, Calame, Nguyen, Le, Bonkowski & Ziegler by having more ventral scale rows (34–42 vs. 27–29) and
fewer precloacal-femoral pores in males (37 vs. 42–43); from C. russelli Bauer by having fewer precloacal pores in
males (8 vs. 15) and fewer dorsal tubercle rows (13–15 vs. 22); from C. saiyok Panitvong, Sumontha, Tunprasert &
Pauwels by its larger size (SVL 71.2–83.2 mm vs. 56.7–61.0 mm), more ventral scale rows (34–42 vs. 23–24),
more precloacal pores in males (8 vs. 5), and fewer dorsal tubercle rows (13–15 vs. 18–19); from C. samroiyot
Pauwels & Sumontha by its larger size (SVL reaching 83.2 mm vs. 66.9 mm), more precloacal pores in males (8 vs.
7), and fewer dorsal tubercle rows (13–15 vs. 17–18); from C. sanook Pauwels, Sumontha, Latinne & Grismer by
having more ventral scale rows (34–42 vs. 27–28), the presence of femoral pores in males (vs. absence), and more
precloacal pores in males (8 vs. 3–4); from C. soni Le, Nguyen, Le & Ziegler by its smaller size (71.2–83.2 mm vs.
88.7–103.0 mm), having more enlarged femoral scales on each thigh (15–17 vs. 8–9), more precloacal-femoral
pores in males (37 vs. 18–22), and the absence of femoral pores in females (vs. 11–14); from C. soudthichaki Luu,
Calame, Nguyen, Bonkowski & Ziegler by having more ventral scale rows (34–42 vs. 32–33) and more precloacal-
femoral pores in males (37 vs. 29); from C. spelaeus Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov,
Konstantinov & Chulisov by its smaller size (71.2–83.2 mm vs. 88.9–91.0 mm), the presence of femoral pores in
males (vs. absence), and fewer lamellae under toe IV (18–21 vs. 22–24); from C. sumonthai Bauer, Pauwels &
Chanhome by the presence of enlarged femoral scales (vs. absence), the presence of femoral pores in males (vs.
absence), having more precloacal pores in males (8 vs. 2), and more lamellae under finger IV (17–19 vs. 16); from
C. surin Chan-ard & Makchai by having more precloacal pores in males (8 vs. 4) and the presence of femoral pores
in males (vs. absence); from C. thochuensis Ngo & Grismer more precloacal pores in males (8 vs. 3–5) and fewer
dorsal tubercle rows (13–15 vs. 20–26); from C. takouensis Ngo & Bauer by having more enlarged femoral scales
on each thigh (15–17 vs. 3–5), more femoral pores on each thigh in males (14–15 vs. 0–2), more precloacal pores
in males (8 vs. 3–4), and generally more lamellae under finger IV (17–19 vs. 16–17); from C. teyniei David,
Nguyen, Schneider & Ziegler by its smaller size (SVL 71.2–83.2 mm vs. 89.9 mm), having fewer enlarged femoral
scales on each thigh (15–17 vs. 23), the presence of femoral pores in males (vs. absence), and a different dorsal
color pattern (banded vs. blotched); from C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome by the presence
of precloacal-femoral pores in males (vs. absence), and having more lamellae under finger IV (17–19 vs. 16); from
C. tigroides Bauer, Sumontha & Pauwels by having more precloacal-femoral pores in males (37 vs. 21) and the
absence of precloacal-femoral pores in females (vs. 21); C. variegatus (Blyth) by having more precloacal-femoral
pores in males (37 vs. 32) and more ventral scale rows (34–42 vs. 22); from C. wangkulangkulae Sumontha,
Pauwels, Suwannakarn, Nutatheera & Sodob by the presence of precloacal-femoral pores in males (vs. absence);
and from C. yangbayensis Ngo & Chan by having more femoral pores on each thigh in males (14–15 vs. 0–2) and
more lamellae under toe IV (18–21 vs. 15–17).
Morphologically, the new species resemble C. huongsonensis and C. soni. However, it can be distinguished
from the latter by having a smaller size and differences in the number of enlarged femoral scales on thighs and the
number of femoral and precloacal pores.
New record of Cyrtodactylus otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, 2015 from Son La
Province (Fig. 6)
Specimens examined (n = 2). One adult male, IEBR A.2017.3 (Field No. SL2016.152) and one adult female, TBU
2017.2 (Field No. SL 2016.151) collected by Nenh Ba Song, 28 June 2016, near Na Bai Village, Chieng Yen
Commune within Xuan Nha NR (20
o
45.711’N, 104
o
56.326’E, elevation 1100 m a.s.l.), Van Ho District, Son La
Province.
In terms of genetic divergence, the sequences of both newly collected samples of C. otai from Van Ho District,
Son La Province were identical to those of the type specimens with GenBank accession numbers KT004370 and
KT004371.
Morphological characters of the two specimens from Son La Province also fit well with the descriptions of
Nguyen et al. (2015a) (see Table 2); SVL 92.5 mm in the male, 88.5 mm in the female; tails of both specimens
regenerated (TaL 61.0 mm in the male, 60.0 mm in the female).
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FIGURE 6. The male (IEBR A.2017.3) of Cyrtodactylus otai in life from Son La Province. Photo N. B. Song.
Body elongate, head distinguished from neck, elongate, depressed; snout long, round anteriorly; snout scales
small, round, granular, larger than those on frontal and parietal regions; eye large, pupils vertical; upper eyelid
fringe with spinous scales; ear oval, small; rostral wider than high with a medial suture; mental triangular, slightly
narrower than rostral; supralabials 10; infralabials 9–10. Dorsal scales granular; dorsal tubercles round, conical,
present on occipital region and back, each surrounded by 8–10 granular scales, in 11–13 irregular longitudinal rows
at midbody; ventral scales smooth, medial scales 2 or 3 times larger than dorsal scales, in 40–41 longitudinal rows
at midbody; lateral skin folds distinct without tubercles; 185–186 ventral scales between mental and cloacal slit;
precloacal groove absent; enlarged femoral scales absent; femoral pores absent; precloacal pores 7 in the male,
absent in the female. Dorsal forelimbs covered by few slightly developed tubercles; dorsal hind limb covered by
distinctly developed tubercles; fingers and toes without webbing; subdigital lamellae of fourth finger 18–19;
subdigital lamellae of fourth toe 19–21. Tail regenerated; postcloacal tubercles 2–3; dorsal tail bearing distinct
tubercles at base; subcaudals not enlarged, flat, smooth.
Coloration in preservative. Head and back greyish cream; dorsal side of head with dark brown markings, in
oval, arched and lozenge shapes on occiput; a dark stripe extending from posterior corner of eye rearwards to above
tympanum; neck with some large dark blotches, forming a discontinuous band; dorsum with five transverse dark
brown bands between fore- and hind-limb insertions, edged in white anteriorly and posteriorly; dorsal surface of
fore and hind limbs with dark blotches and bars; tail greyish cream; chin, throat, chest, belly and lower limbs
pinkish white. For coloration in life see Fig. 6.
Ecological notes. Two specimens were found between 19:20 and 21:05h, on tree branches, near limestone
cliffs. The surrounding habitat was disturbed evergreen forest of medium and small hardwood and shrub.
Distribution. C. otai was recently described from Hang Kia – Pa Co Nature Reserve in Hoa Binh Province
(Nguyen et al. 2015a). The new recorded locality from Son La Province is approximately 30 km NW from the type
locality (Fig. 1).
Discussion
It is intriguing that Bayesian and ML analyses support the sister relationship between C. sonlaensis and C. cf.
pulchellus, although previous studies, e.g., Nguyen et al. (2015a), Le et al. (2016), Luu et al. (2016b), and the MP
analysis in this study recovered a more basal position of the latter taxon. This relationship could be an artifact of
long-branch attraction due to either incomplete taxon sampling or rate heterogeneity in the COI gene (Bergsten
2005). As a result of using a short fragment of the COI gene, the placements of other taxa of the genus
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A NEW CYRTODACTYLUS FROM VIETNAM
Cyrtodactylus in the study were either weakly corroborated or unresolved (Fig. 1), suggesting the necessity of
adding more independent molecular markers, especially slow-evolving nuclear genes. We are currently
investigating phylogenetic relationships of Cyrtodactylus species in Vietnam and neighboring countries using more
mitochondrial and nuclear molecular markers. The results are expected to shed light on these issues.
The discovery of a new species in limestone karsts further highlights the importance of these ecosystems in
generating and harboring endemic diversity. Until recently, vertebrate fauna had been poorly studied in Southeast
Asian karsts (Clements et al. 2006). However, the latest works uncovered a high level of microendemism across a
wide variety of taxonomic groups in these unique ecosystems (Chung et al. 2014; Grismer et al. 2016; Luu et al.
2016a; Nicolas et al. 2012). Although northern Vietnam has one of the most extensive limestone karst systems in
the region (Clements et al. 2006), it remains insufficiently understood. Recent surveys, e.g., Ngo & Chan (2011),
Nguyen et al. (2015a), Grismer et al. (2016), and Le et al. (2016), continue to discover new species of
Cyrtodactylus from the region. However, it is likely that additional new species in the genus will be found if more
expeditions are conducted in the karst system.
Acknowledgements
We thank N.V. Cam and H.V. Dinh (Muong Bang Commune), H.V. Tu and N.B. Song (Tay Bac University), T.Q.L.
Hoang (Binh Thuan High School) for their assistance in the field. We thank E. Sterling (New York) and K. Koy
(Berkeley) for providing the original map. We are grateful to L.L. Grismer (Riverside, California), O.G.S. Pauwels
(Brussels), and A. Bauer (Villanova) for their helpful comments. This research is supported by the National
Foundation for Science and Technology Development (NAFOSTED, Grant No. 106-NN.06-2016.59) and the
National Geographic (Grant No. 230151).
References
Bauer, A.M. (2002) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from Myanmar. Proceedings of the California
Academy of Sciences, 53 (7), 75–88.
Bauer, A.M. (2003) Descriptions of seven new Cyrtodactylus (Squamata: Gekkonidae) with a key to the species of Myanmar
(Burma). Proceedings of the California Academy of Sciences, 54 (25), 463–498.
Bauer, A.M., Kunya, K., Sumontha, M., Niyomwan, P., Panitvong, N., Pauwels, O.S.G., Chanhome, L. & Kunya, T. (2009)
Cyrtodactylus erythrops (Squamata: Gekkonidae), a new cave-dwelling gecko from Mae Hong Son Province, Thailand.
Zootaxa, 2124, 51–62.
Bauer, A., Kunya, K., Sumontha, M., Niyomwan, P., Pauwels, O.S.G., Chanhome, L. & Kunya, T. (2010) Cyrtodactylus
dumnuii (Squamata: Gekkonidae), a new cave–dwelling gecko from Chiang Mai Province, Thailand. Zootaxa, 2570, 41–
50.
Bauer, A.M., Pauwels, O.S.G. & Chanhome, L. (2002) A new species of cave-dwelling Cyrtodactylus (Squamata: Gekkonidae)
from Thailand. The Natural History Journal of Chulalongkorn University, 2, 19–29.
Bauer, A.M., Sumontha, M. & Pauwels, O.S.G. (2003) Two new species of Cyrtodactylus (Reptilia: Squamata: Gekkonidae)
from Thailand. Zootaxa, 376, 1–18.
https://doi.org/10.11646/zootaxa.376.1.1
Bergsten, J. (2005) A review of long-branch attraction. Cladistics, 21, 163–193.
https://doi.org/10.1111/j.1096-0031.2005.00059.x
Chan-ard, T. & Makchai, S. (2011) A new insular species of Cyrtodactylus Gray, 1827 (Squamata, Gekkonidae), from the Surin
Islands, Phang-nga Province, southern Thailand. The Thailand Natural History Museum Journal, 5 (1), 7–15.
Chung, K.-F., Leong, W.-C., Rubite, R.R., Repin, R., Kiew, R., Liu, Y. & Peng, C.-I. (2014) Phylogenetic analyses of Begonia
sect. Coelocentrum and allied limestone species of China shed light on the evolution of Sino-Vietnamese karst flora.
Botanical Studies, 55, 1
Clements, R., Sodhi, N.S., Schilthuizen, M. & Ng, P.K.L. (2006) Limestone karsts of Southeast Asia: imperiled arks of
biodiversity. BioScience, 56, 733–742.
https://doi.org/10.1641/0006-3568(2006)56[733:LKOSAI]2.0.CO;2
Connette, G.M., Oswald, P., Thura, M.K., Connette, K.J.L., Grindley, M.E., Songer, M., Zug, R.G. & Mulcahy, G.D. (2017)
Rapid forest clearing in a Myanmar proposed national park threatens two newly discovered species of geckos
(Gekkonidae: Cyrtodactylus). PLOS ONE, 12 (4), e0174432.
https://doi.org/10.1371/journal.pone.0174432
David, P., Nguyen, T.Q., Schneider, N. & Ziegler, T. (2011) A new species of the genus Cyrtodactylus Gray, 1827 from central
Laos (Squamata: Gekkonidae). Zootaxa, 2833, 29–40.
NGUYEN ET AL.
38
·
Zootaxa 4341 (1) © 2017 Magnolia Press
Grismer, L.L. & Ahmad, N. (2008) A new insular species of Cyrtodactylus (Squamata: Gekkonidae) from the Langkawi
Archipelago, Kedah, Peninsular Malaysia. Zootaxa, 1924, 53–68.
Grismer, L.L., Wood, P.L. Jr., Anuar, S., Davis, H.R., Burch, B.T., Cobos, A.J. & Murdoch, M.L. (2016) A new species of karst
forest Bent-toed Gecko (genus Cyrtodactylus Gray) not yet threatened by foreign cement companies and a summary of
Peninsular Malaysia’s endemic karst forest herpetofauna and the need for its conservation, Zootaxa, 4061 (1), 1–17.
https://doi.org/10.11646/zootaxa.4061.1.1
Grismer, L.L., Wood, P.L. Jr., Quah, E.S.H., Anuar, S., Muin, A., Sumontha, M., Ahmad, N., Bauer, A.M., Wangkulangkul, S.,
Grismer, J.L. & Pauwels, O.S.G. (2012) A phylogeny and taxonomy of the Thai-Malay Peninsula Bent-toed Geckos of the
Cyrtodactylus pulchellus complex (Squamata: Gekkonidae): combined morphological and molecular analyses with
descriptions of seven new species. Zootaxa, 3520, 1–55.
Hoang, Q.X., Orlov, N.L., Ananjeva, N.B., Johns, A.G., Hoang, T.N. & Dau, V.Q. (2007) Description of a new species of the
genus Cyrtodactylus Gray, 1827 (Squamata: Sauria: Gekkonidae) from the karst of North Central Vietnam. Russian
Journal of Herpetology, 14, 98–106.
Ivanova, N.V., de Waard, J. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality
DNA. Molecular Ecology Notes, 6, 998–1002.
https://doi.org/10.1111/j.1471-8286.2006.01428.x
Kunya, K., Panmongkol, A., Pauwels, O.G.S., Sumontha, M., Meewasana, J., Bunkhwamdi, W. & Dangsri, S. (2014) A new
forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Doi Suthep, Chiang Mai Province,
northern Thailand. Zootaxa, 3811 (2), 251–261.
https://doi.org/10.11646/zootaxa.3811.2.6
Kunya, K., Sumontha, M., Panitvong, N., Dongkumfu, W., Sirisamphan, T. & Pauwels, O.S.G. (2015) A new forest-dwelling
Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Doi Inthanon, Chiang Mai Province, northern Thailand.
Zootaxa, 3905 (4), 573–584.
https://doi.org/10.11646/zootaxa.3905.4.9
Le, D.T., Nguyen, T.Q., Le, M.D. & Ziegler, T. (2016) A new species of Cyrtodactylus (Squamata: Gekkonidae) from Ninh
Binh Province, Vietnam. Zootaxa, 4162 (2), 268–282.
https://doi.org/10.11646/ zootaxa.4162.2.4
Le, T.D., Nguyen, T.T., Nishikawa, K., Nguyen, L.H.S., Pham, V.A., Matsui, M., Bernardes, M., & Nguyen, Q.T. (2015a) A
New Species of Tylototriton Anderson, 1871 (Amphibia: Salamandridae) from Northern Indochina. Current Herpetology,
34, 38–50.
Le, T.D., Pham, V.A., Nguyen, L.H.S., Ziegler, T. & Nguyen, Q.T. (2014) Babina lini (Chou, 1999) and Hylarana menglaensis
Fei, Ye, and Xie, 2008, two additional anuran species for the herpetofauna of Vietnam. Russian Journal of Herpetology,
21, 315–321.
Le, T.D., Pham, V.A., Nguyen, L.H.S., Ziegler, T. & Nguyen, Q.T. (2015b) First records of Megophrys daweimontis Rao and
Yang, 1997 and Amolops vitreus (Bain, Stuart and Orlov, 2006) (Anura: Megophryidae, Ranidae) from Vietnam. Asian
Herpetological Research, 6, 66–72.
Le, T.D., Pham, V.A., Pham, T.C., Nguyen, L.H.S., Ziegler, T. & Nguyen, Q.T. (2015c) Review of the genus Sinonatrix in
Vietnam with a new country record of Sinonatrix yunnanensis Rao and Yang, 1998. Russian Journal of Herpetology, 22,
84–88.
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.
https://doi.org/10.1016/j.ympev.2006.03.003
Luu, V.Q., Bonkowski, M., Nguyen, T.Q., Le, M.D., Schneider, N., Ngo, H.T. & Ziegler, T. (2016a) Evolution in karst massifs:
Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new
country record from Laos. Zootaxa, 4107 (2), 101–140.
https://doi.org/10.11646/zootaxa.4107.2.1
Luu, V.Q., Calame, T., Bonkowski, M., Nguyen, T.Q. & Ziegler, T. (2014) A new species of Cyrtodactylus (Squamata:
Gekkonidae) from Khammouane Province, Laos. Zootaxa, 3760 (1), 54–66.
https://doi.org/10.11646/zootaxa.3760.1.3
Luu, V.Q., Calame, T., Nguyen, T.Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata:
Gekkonidae) from the limestone forest of Khammouane Province, central Laos. Zootaxa, 4058 (3), 388–402.
https://doi.org/10.11646/zootaxa.4058.3.6
Luu, V.Q., Calame, T., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016b) Cyrtodactylus rufford, a new cave-
dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2), 185–
199.
https://doi.org/10.11646/zootaxa.4067.2.4
Luu, V.Q., Nguyen, T.Q., Do, H.Q. & Ziegler, T. (2011) A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son
limestone forest, Hanoi, northern Vietnam. Zootaxa, 3129, 39–50.
Luu, V.Q., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016c) A new species of karst-dwelling Bent-toed Gecko
(Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 87–102.
https://doi.org/10.11646/zootaxa.4079.1.6
Nazarov, R.A., Poyarkov, N.A., Orlov, N.L., Nguyen, S.N., Milto, K.D., Martynov, A.A., Konstantinov, E.L. & Chulisov, A.S.
(2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new
species. Proceedings of the Zoological Institute RAS, 318, 391–423.
Zootaxa 4341 (1) © 2017 Magnolia Press
·
39
A NEW CYRTODACTYLUS FROM VIETNAM
Ngo, T.V. (2008) Two new cave-dwelling species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Southwestern Vietnam.
Zootaxa, 1909, 37–51.
Ngo, T.V. & Bauer, A.M. (2008) Descriptions of two new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae)
endemic to southern Vietnam. Zootaxa, 715, 27–42.
Ngo, T.V. & Chan, K.O. (2010) A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa
province, southern Vietnam. Zootaxa, 2504, 47–60.
Ngo, T.V. & Chan, K.O. (2011) A new karstic cave-dwelling Cyrtodactylus Gray (Squamata: Gekkonidae) from northern
Vietnam . Zootaxa, 3125, 51–63.
Ngo, T.V. & Grismer, L.L. (2010) A new karst dwelling Cyrtodactylus (Squamata: Gekkonidae) from Son La Province,
northwestern Vietnam. Hamadryad, 35, 84–95.
Ngo, V.T. & Grismer, L.L. (2012) A new endemic species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu
Island, southwestern Vietnam. Zootaxa, 3228 (1), 48–60.
https://doi.org/10.11646/zootaxa.3228.1.2
Ngo, V.T., Grismer, L.L. & Grismer, J.L. (2008) A new endemic cave dwelling species of Cyrtodactylus Gray, 1827 (Squamata:
Gekkonidae) in Kien Giang Biosphere Reserve, southwestern Vietnam. Zootaxa, 1967, 53–62.
Ngo, V.T., Grismer, L.L. & Grismer, J.L. (2010) A new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) in Phu
Quoc National Park, Kien Giang Biosphere Reserve, southwestern Vietnam. Zootaxa, 2604, 37–51.
Ngo, T.V. & Pauwels, O.S.G. (2010) A new cave-dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from
Khammouane Province, southern Laos. Zootaxa, 2730 (1), 44–56.
https://doi.org/10.11646/zootaxa.2730.1.3
Nguyen, S.N., Orlov, N.L. & Darevsky, S.I. (2006) Descriptions of two new species of the genus Cyrtodactylus Gray, 1827
(Squamata: Sauria: Gekkonidae) from Southern Vietnam. Russian Journal of Herpetology, 13, 215–226.
Nguyen, S.N., Yang, J-X., Le, N.T.T., Nguyen, L.T., Orlov, N.L., Hoang, C.V., Nguyen, T.Q., Jin, J-Q., Rao, D-Q., Hoang, T.N.,
Che, J., Murphy, R.W. & Zhang, Y-P. (2014) DNA barcoding of Vietnamese bent-toed geckos (Squamata: Gekkonidae:
Cyrtodactylus) and the description of a new species. Zootaxa, 3784 (1), 48–66.
https://doi.org/10.11646/zootaxa.3784.1.2
Nguyen, T.Q., Le, M.D., Pham, A.V., Ngo, H.N., Hoang, C.V., Pham, C.T. & Ziegler, T. (2015a) Two new species of
Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of Hoa Binh Province, Vietnam. Zootaxa, 3985 (3), 375–390.
https://doi.org/10.11646/zootaxa.3985.3.3
Nguyen, T.Q, Pham, V.A., Nguyen, L.H.S., Le, D.M. & Ziegler, T. (2015b) First record of Parafimbrios lao Teynié, David,
Lottier, Le, Vidal & Nguyen, 2015 (Squamata: Xenodermatidae) from Vietnam. Russian Journal of Herpetology, 22, 297–
300.
Nicolas, V., Herbreteau, V., Couloux, A., Keovichit, K., Douangboupha, B. & Hugot, J.-P. (2012) A remarkable case of micro-
endemism in Laonastes aenigmamus (Diatomyidae, Rodentia) revealed by nuclear and mitochondrial DNA sequence data.
PLOS ONE, 7, e48145.
https://doi.org/10.1371/journal.pone.0048145
Orlov, N.L., Nguyen, Nazarov, R.A., Anajeva, N.B. & Nguyen, S.N. (2007) A new species of the genus Cyrtodactylus Gray,
1827 and redescription of Cyrtodactylus paradoxus (Darevsky et Szczerbak, 1997) [Squamata: Sauria: Gekkonidae] from
south Vietnam. Russian Journal of Herpetology, 14, 145–152.
Panitvong, N., Sumontha, M., Tunprasert, J. & Pauwels, O.S.G. (2014) Cyrtodactylus saiyok sp. nov., a new dry evergreen
forest-dwelling Bent-toed Gecko (Squamata: Gekkonidae) from Kanchanaburi Province, western Thailand. Zootaxa, 3869
(1), 64–74.
https://doi.org/10.11646/zootaxa.3869.1.6
Pauwels, O.S.G., Bauer, A.M., Sumontha, M. & Chanhome, L. (2004) Cyrtodactylus thirakhupti (Squamata: Gekkonidae), a
new cave-dwelling gecko from southern Thailand. Zootaxa, 772, 1–11.
Pauwels, O.S.G. & Sumontha, M. (2014) Cyrtodactylus samroiyot, a new limestone-dwelling Bent-toed Gecko (Squamata:
Gekkonidae) from Prachuap Khiri Khan Province, peninsular Thailand. Zootaxa, 3755 (6), 573–583.
https://doi.org/10.11646/zootaxa.3755.6.4
Pauwels, O.S.G., Sumontha, M. & Bauer, A.M. (2016) A new Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from
Phetchaburi Province, Thailand. Zootaxa, 4088 (3), 409–419.
https://doi.org/10.11646/zootaxa.4088.3.6
Pauwels, O.S.G., Sumontha, M., Keeratikiat, K. & Phanamphon, E. (2014a) Cyrtodactylus kunyai (Squamata: Gekkonidae), a
new cave-dwelling Bent-toed Gecko from Loei Province, northeastern Thailand. Zootaxa, 3821 (2), 253–264.
Pauwels, O.S.G., Sumontha, M., Latinne, A. & Grismer, L.L. (2013) Cyrtodactylus sanook (Squamata: Gekkonidae), a new
cave-dwelling gecko from Chumphon Province, southern Thailand. Zootaxa, 3635 (3), 275–285.
https://doi.org/10.11646/zootaxa.3635.3.7
Pauwels, O.S.G., Sumontha, M., Panitvong, N. & Varaguttanonda, V. (2014b) Cyrtodactylus khelangensis, a new cave-dwelling
Bent-toed Gecko (Squamata: Gekkonidae) from Lampang Province, northern Thailand. Zootaxa, 3755 (6), 584–594.
Pham, V.A., Le, T.D., Nguyen, L.H.S., Ziegler, T. & Nguyen, Q.T. (2014) First records of Leptolalax eos Ohler, Wollenberg,
Grosjean, Hendrix, Vences, Ziegler & Dubois, 2011 and Hylarana cubitalis (Smith, 1917) (Anura: Megophryidae,
Ranidae) from Viet Nam. Russian Journal of Herpetology, 21, 195–200.
Pham, V.A., Le, T.D., Pham, T.C., Nguyen, L.H.S., Ziegler, T. & Nguyen, Q.T. (2016) Two additional records of megophryid
species, Leptolalax minimus (Taylor, 1962) and Leptobrachium masatakasatoi Matsui, 2013, for the herpetofauna of
NGUYEN ET AL.
40
·
Zootaxa 4341 (1) © 2017 Magnolia Press
Vietnam . Revue suisse de Zoologie, 123, 35–43.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
https://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.
Schneider, N., Nguyen, T.Q., Schmitz, A., Kingsada, P., Auer, M. & Ziegler, T. (2011) A new species of karst dwelling
Cyrtodactylus (Squamata: Gekkonidae) from northwestern Laos. Zootaxa, 2930, 1–21.
Simmons, J.E. (2002) Herpetological collecting and collections management. Revised edition. Society for the Study of
Amphibians and Reptiles. Herpetological Circular, 31, 1-153.
Smith, M.A. (1917) Descriptions of new reptiles and a new batrachian from Siam. Journal of Natural History Society of Siam,
2, 221–225.
Smith, M.A. (1921a) New or little-known reptiles and batrachians from southern Annam (Indo-China). Proceedings of the
Zoological Society of London, 1921, 423–440.
Smith, M.A. (1921b) Reptiles and Batrachians collected on Pulo Condore. Journal Natural History Society, 4, 93–97. [1920]
Smith, M.A. (1935) The fauna of British India, including Ceylon and Burma. Vol. II. Sauria. Taylor and Francis, London, 8 +
440 pp.
Sumontha, M., Panitvong, N. & Deein, G. (2010) Cyrtodactylus auribalteatus (Squamata: Gekkonidae), a new cave-dwelling
gecko from Phitsanulok Province, Thailand. Zootaxa, 2370, 53–64.
Sumontha, M., Pauwels, O.S.G., Kunya, K., Nitikul, A., Samphanthamit, P. & Grismer, L.L. (2012) A new forest-dwelling
gecko from Phuket Island, Southern Thailand, related to Cyrtodactylus macrotuberculatus (Squamata: Gekkonidae).
Zootaxa, 3522, 61–72.
Sumontha, M., Pauwels, O.S.G., Suwannakarn, N., Nutatheera, N. & Sodob, W. (2014) Cyrtodactylus wangkulangkulae
(Squamata: Gekkonidae), a new Bent-toed Gecko from Satun Province, southern Thailand. Zootaxa, 3821 (1), 116–124.
https://doi.org/10.11646/zootaxa.3821.1.8
Swofford, D.L. (2001) PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4. Sinauer Associates,
Sunderland, Massachusetts.
Taylor, E.H. (1963) Lizards of Thailand. The Universtiy of Kansas Science Bulletin, 55, 687–1077.
The People's Committee of Son La Province (2007) Son La Province Portal. Available from: http://sonla.gov.vn/gioi-thieu
(accessed 25 March 2017)
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: Xexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876–4882.
Ulber, T. (1993) Bemerkungen über cyrtodactyline Geckos aus Thailand nebst Beschreibungen von zwei neuen Arten (Reptilia:
Gekkonidae). Mitteilungen aus dem Zoologischen Museum in Berlin, 69, 187–200.
https://doi.org/10.1002/mmnz.19930690202
Ziegler, T., Nazarov, R., Orlov, N., Nguyen, T.Q., Vu, T.N., Dang, K.N., Dinh, T.H. & Schmitz, A. (2010) A third new
Cyrtodactylus (Squamata: Gekkonidae) from Phong Nha – Ke Bang National Park, Vietnam. Zootaxa, 2413, 20–36.
Ziegler, T., Phung, T.M., Le, M.D. & Nguyen, T.Q. (2013) A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen
Province, southern Vietnam. Zootaxa, 3686 (4), 432–446.
https://doi.org/10.11646/zootaxa.3686.4.2
Ziegler, T., Rösler, H., Herrmann, H.-W. & Vu, T.N. (2002) Cyrtodactylus phongnhakebangensis sp. n., ein neuer
Bogenfingergecko aus dem annamitischen Karstwaldmassiv, Vietnam. Herpetofauna, 24, 11–25.
APPENDIX. Comparative specimens examined
Cyrtodactylus bichnganae. Vietnam: Son La: Son La City: TBU PAT.250, TBU NT.2014.01.
C. bobrovi. Vietnam: Hoa Binh: Ngoc Son – Ngo Luong: IEBR A.2015.29 (holotype), IEBR A.2015.30, VNMN A.2015.61,
VFM 2015.1 (paratypes).
C. jaegeri. Laos: Khammouane: Thakhek: IEBR A.2013.55 (holotype), NUOL R-2013.1 (paratype), VFU TK.914.
C. huongsonensis. Vietnam: Hanoi: Huong Son: IEBR A.2011.3 (holotype), ZFMK 92293 (paratype).
C. cf. lomyenensis. Laos: Khammouane: Phou Hin Boun: IEBR KM2012.52, KM2012.54, KM2012.57.
C. otai. Vietnam: Hoa Binh: Hang Kia – Pa Co: IEBR A.2015.26 (holotype), IEBR A.2015.27–28, VNMN A.2015.60, ZFMK
96721 (paratypes).
C. pageli. Laos: Vientiane Province: Vang Vieng: IEBR A.2010.36 (holotype), IEBR A.2010.37, MTD 48025, MHNG
2723.91, NUOL 2010.3–2010.7, ZFMK 91827 (paratypes).
C. roesleri. Vietnam: Quang Binh Province: Phong Nha – Ke Bang: ZFMK 89377 (holotype), IEBR A.0932, MHNG 2713.79,
VNUH 220509, ZFMK 86433, 89378 (paratypes).
C. teyniei. Laos: Borikhamxay Province: near Ban Na Hin: NEM 0095 (holotype); Khammouane Province: Ban Na Than:
IEBR KM.2012.14–15.
C. vilaphongi. Laos: Luang Prabang: IEBR A.2013.103 (holotype), NUOL R-2013.5 (paratype).
C. wayakonei. Laos: Luang Nam Tha: Vieng Phoukha: IEBR A.2010.01 (holotype), ZFMK 91016, MTD 47731, NUOL 2010.1
(paratypes).
... Sequences of C. cf. interdigitalis Ulber, 1993 and C. elok Dring, 1979 were used as outgroups according to Nguyen et al. (2017) and Schneider et al. (2020). ...
... Bilateral scale counts were given as left/right. The methodology of measurements and meristic counts followed Ngo (2011) Morphological comparisons and analyses were based on specimen examination and data obtained from the literature (Hoang et al. 2007;Rösler et al. 2008;Bauer et al. 2009Bauer et al. , 2010Ngo and Grismer 2010;Nguyen et al. 2010Nguyen et al. , 2015Nguyen et al. , 2017Sumontha et al. 2010;Teynié and David 2010;Luu et al. 2011Luu et al. , 2013Luu et al. , 2016Ngo 2011;Ngo and Chan 2011;Schneider et al. 2011Schneider et al. , 2014Schneider et al. , 2020Kunya et al. 2014;Nazarov et al. 2014Nazarov et al. , 2018Nguyen et al. 2014;Pham et al. 2019). ...
... The obtained sequence alignment is 690 bp in length. The topologies derived from ML and BI analyses were similar and basically consistent with those of Nguyen et al. (2017), Pham et al. (2019), andSchneider et al. (2020). The sequences of three specimens collected from Zhenkang County, Yunnan, China were nested them within the Cyrtodactylus wayakonei group and the sister group to a clade consisting of C. wayakonei Nguyen, Kingsada, Rösler, Auer &Ziegler, 2010 andC. ...
Article
Full-text available
A new species of Cyrtodactylus is described on the basis of five specimens collected from the karst formations of Zhenkang County, Yunnan Province, China. Cyrtodactylus zhenkangensis sp. nov. is recognized by having a unique combination of morphological characters, the most diagnostic being: 12–15 enlarged femoral scales on each thigh; 2–5 femoral pores on each thigh in males, 0–3 pitted scales on each thigh in females; eight or nine precloacal pores in a continuous row or separated by one poreless scale in males, 7–9 pitted scales in females; subcaudals enlarged, arranged alternately as single and double on anterior and mostly single at middle and posterior; dorsal surface of head with obvious reticulations. Phylogenetic analyses show that the new species is a member of the C. wayakonei species group and a sister taxon to a clade consisting of C. wayakonei and C. martini based on Maximum Likelihood analyses and Bayesian Inference and differs from its congeners by at least 12.0% genetic divergence in a fragment of the COI gene.
... At present, the genus contains more than 300 recognized species (Uetz et al., 2021), and is widely distributed across South Asia to Melanesia where they occupy a vast territory (Termprayoon et al., 2021). Le et al. (2016) designated the Cyrtodactylus wayakonei species group in the genus Cyrtodactylus containing ten species, subsequently, Nguyen et al. (2017), Pham et al. (2019), Schneider et al. (2020), and Liu and Rao (2021) expanded this group to include 17 species. Gris-mer et al. (2021b) repartitioned the species group of Cyrtodactylus and revised C. wayakonei species group to C. chauquangensis species group. ...
... The methodology of measurements and meristic counts followed Liu and Rao (2021): snout-vent length (SVL), from tip of snout to anterior margin of cloaca; tail length (TaL), from posterior margin of cloaca to tip of tail; maximum head height (HH), from occiput to underside of jaws; head length (HL), from tip of snout to posterior margin of ear; maximum head width (HW); greatest diameter of orbit (OD); snout to eye distance (SE), from tip of snout to anterior corner of eye orbit; eye orbit to ear distance (EE), from posterior corner of eye orbit to anterior margin of ear opening; internarial distance (IND), measured between inner borders of nostrils; interorbital distance (IOD), measured across narrowest point of frontal bone; ear diameter (ED), greatest diameter of ear; axilla to groin distance (AG); forearm length (ForeaL), from the base of the palm to the elbow; shank length (SL), from the base of heel to the knee; rostral width (RW); rostral height (RH); mental width (MW); mental length (ML); supralabials (SPL); infralabials (IFL); internasals or postrostrals (I); postmentals (PM), i.e., scales bordering mental shield, except infralabials; granular scales surrounding dorsal midbody tubercles (GSDT); dorsal tubercle rows (DTR), number of dorsal, longitudinal rows of tubercles at midbody between the ventrolateral folds; paravertebral tubercles (PVT), counted in a single paravertebral row from the level of the forelimb insertions to the level of the hind limb insertion; longitudinal ventral scale rows (V), counted across the belly between the ventrolateral folds at midbody; enlarged femoral scales (EFS), number of enlarged femoral scale beneath each thigh; precloacal pores (PP); femoral pores (FP); postcloacal tubercles (PAT), number of tubercles on each side of postcloacal region; subdigital lamellae under the fourth finger (LD4); subdigital lamellae under the fourth toe (LT4). Morphological comparisons were based on the original descriptions of each species (Hoang et al., 2007;Bauer et al., 2009Bauer et al., , 2010Ngo and Grismer, 2010;Nguyen et al., 2010Nguyen et al., , 2017Sumontha et al., 2010;Luu et al., 2011;Ngo 2011;Ngo and Chan, 2011;Kunya et al., 2014;Nazarov et al., 2014;Nguyen et al., 2014;Schneider et al., 2014Schneider et al., , 2020Le, 2016;Pham et al., 2019;Liu and Rao, 2021). Molecular analyses. ...
Article
A new species of Cyrtodactylus is described based on five specimens collected from the karst formations of Maguan County, Wenshan Prefecture, Yunnan Province, China. The new species is recognized by having a unique combination of morphological characters: medium body size, ventrolateral folds present without interspersed tubercles, 7 – 9 precloacal pores in a continuous series, enlarged femoral scales present and continuous with pore-bearing precloacal scales, femoral pores on each enlarged femoral scale in males, 1 – 4 postcloacal tubercles on each side, subcaudals enlarged, a black postocular streak extending from posterior corner of eye rearwards to above tympanum, nuchal loop discontinuous, 6 – 7 black irregular dorsal bands between limbs, most bands discontinuous. Genetically, uncorrected sequence divergences of the ND2 gene and its flanking tRNAs between the new species and investigated congeners ranged from 12.5% to 18.2%.
... Considering our samples were collected near Vietnam and Laos, 40 COI sequences of Cyrtodactylus species (and one sequence of Cyrtopodion for our outgroup) from previous studies (Nguyen et al., , 2017Ziegler et al., 2013;Schendier et al., 2014;Le et al., 2016;Luu et al., 2016aLuu et al., , 2016bBrennan et al., 2017;Connette et al., 2017; from Southeast Asian specimens were downloaded from Genbank (National Center for Biotechnology Information) to explore the relationships with other known species (Table 1). All 46 COI sequences were initially aligned by using Geneious Basic (Kearse et al., 2012), and then optimized by MEGA X (Kumar et al., 2018). ...
... nov. can be distinguished from C. sonlaensis(Nguyen, Pham, Ziegler, Ngo and Le, 2017) by having Morphological measurements and scalation from holotype and paratypes of Cyrtodactylus hekouensis sp. nov. ...
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Cyrtodactylus geckos are one of the most speciose and diverse groups of extant lizards known, distributed throughout the Asian and Pacific realms. Using molecular phylogenetic methods and supporting morphological data, we describe a new species of Cyrtodactylus in Daweishan National Nature Reserve, Yunnan Province, China. Cyrtodactylus hekouensis sp. nov. can be morphologically distinguished from its nearby congeners by the following characters: maximum SVL 92.3 mm and TL 98.5 mm; 11-12 supralabials; 11-12 infralabials; 36-57 scale rows between the fifth supralabials; 10-13 dorsal tubercles rows; 3 postnasals on blunt and smooth front snout; precloacal-femoral pores in a continuous series of 33-39 (females with pitted scales) located under vent/cloaca and thighs in both sexes; precloacal groove absent; 3/3 postcloacal tubercles; subdigital lamellae under the fourth finger 21 or 22, under the fourth toe 20-23; smooth midbody with smooth venter and tuberculate dorsal scale rows, tubercles from head to tail base; dorsal transverse patterns are generally large, bilaterally symmetrical. The results of the phylogenetic analysis recover specimens of this new species as sister to a clade containing C. wayakonei and C. martini. Uncorrected pairwise intraspecific distances were < 1%, and distances between our new species and other Cyrtodactylus species from nearby countries ranged from 14.2% to 26.8%.
... Tail longer than SVL (TaL 74.1 mm); 3/2 postcloacal tubercles; dorsal surface of tail bearing distinct tubercles on anterior one-half of tail; lateral rows of spinose scales present; median subcaudal scales slightly enlarged, flat, smooth. Nazarov et al. 2014;Nguyen et al. 2014;Panitvong et al. 2014;Schneider et al. 2014a, b;Nurngsomsri et al. 2014;Grismer et al. 2015;Sumontha et al. 2015;Luu et al. 2015;Luu et al. 2016a,b,c;Le et al. 2016;Luu et al. 2017;Nguyen et al. 2017;Pauwels et al. 2018;Chuaynkern et al. 2018;Nazarov et al. 2018;Murdoch et al. 2019;Pham et al. 2019;Sitthivong et al. 2019;Schneider et al. 2020;Ostrowski et al. 2020Ostrowski et al. , 2021. Coloration in life. ...
... nov. with other congeners from Vietnam and neighboring countries in the mainland Indochina region, including Laos, Cambodia, and Thailand based on examination of specimens (see Appendix) and data from literature (Luu et al. 2014;Nazarov et al. 2014;Nguyen et al. 2014;Panitvong et al. 2014;Schneider et al. 2014 a,b;Nurngsomsri et al. 2014;Grismer et al. 2015;Luu et al. 2015;Sumontha et al. 2015;Pauwels et al. 2016;Le et al. 2016;Luu et al. 2016aLuu et al. ,b,c, 2017Nguyen et al. 2017;Chuaynkern et al. 2018;Pauwels et al. 2018;Nazarov et al. 2018;Murdoch et al. 2019;Pham et al. 2019;Sitthivong et al. 2019;Schneider et al. 2020;Ostrowski et al. 2020Ostrowski et al. , 2021. The new species can be differentiated from other known species of the genus Cyrtodactylus by morphological characters (see Table 3). ...
Article
A new species of the genus Cyrtodactylus is described from Dien Bien Province, northwestern Vietnam based on morphological and molecular data. Cyrtodactylus ngati sp. nov. can be distinguished from remaining congeners by the following combination of characters: maximum SVL 69.3 mm; dorsal pattern consisting of six dark irregular transverse bands between limb insertions; inter-supranasals one; dorsal tubercles present on occiput, body, hind limbs and on first half of tail; 17-22 irregular dorsal tubercle rows at midbody; lateral folds clearly defined, with interspersed tubercles; 32-38 ventral scales between ventrolateral folds; 13 precloacal pores separated by a diastema of 5/5 poreless scales from a series of 7/7 femoral pores in enlarged femoral scales; precloacal and femoral pores absent in females; 1-3 postcloacal tubercles on each side; transversely enlarged median subcaudal scales absent. In the molecular analyses, the new species is shown to be the sister taxon to C. interdigitalis from Thailand. This is the 47 th species of the genus Cyrtodactylus and the first member of the C. brevipalmatus species group recorded from Vietnam.
... In the ML analysis, the -Ln likelihood score of the best tree found was 3920.996. The topology derived from the BA was similar to those of Nguyen et al. (2017) and Brennan et al. (2017). One new species was strongly supported as a sister taxon of Cyrtodactylus puhuensis, while the other one was placed at the basal position of the clade consisting of all species of the C. wayakonei species group, except for C. cf. ...
... In terms of genetic divergence, the new taxon from Houaphan Province is separated by approximately 3.3% from its sister species, C. puhuensis, and the taxon from Luang Prabang Province is divergent from other related taxa by at least 11.6% based on the COI fragment. Pairwise genetic distance between other species in the group is provided in Nguyen et al. (2017). Bayesian cladogram based on the partial COI gene. ...
Article
Two new Cyrtodactylus species are described from Houaphan and Luang Prabang provinces in Laos based on morphological and molecular data. Cyrtodactylus houaphanensis sp. nov. differs from all other Cyrtodactylus in the C. wayakonei species group by at least 3.3 % genetic divergence in the COI gene and can be diagnosed in morphology as follows: SVL 75.8 mm; supralabials 9 or 10; infralabials 8 or 9; ventral scales 35; dorsal tubercles in 20 rows at midbody; precloacal pores 6 in the male; femoral pores absent; subcaudals enlarged; five irregular, brown bands between limb insertions. The new species morphologically resembles C. chauquangensis and revealed to be a sister taxon to C. puhuensis according to our genetic analyses, from which it mainly differs in the absence of enlarged femoral scales. Cyrtodactylus ngoiensis sp. nov. differs from other closely related congeners by at least 11.6 % genetic divergence in the COI gene and can be diagnosed in morphology as follows: maximum SVL 95.3 mm; supralabials 6-9; infralabials 8-11; ventral scales 38-43; dorsal tubercles in 15-21 rows at midbody; enlarged femoral scales present; precloacal pores 7 in the male, 7 pitted scales in females; femoral pores 14 in the male, absent in females; five bright yellowish transverse dorsal bands; subcaudals enlarged. The new species is shown to be a member of the C. wayakonei species group, but morphologically resembles C. dumnuii from Thailand. According to our phylogenetic analyses, it is a basal taxon to a clade comprising C. spelaeus, C. chauquangensis, C. vilaphongi, C. cucphuongensis, C. puhuensis, C. houaphanensis, C. otai and C. bobrovi. Based on a new record of C. bansocensis from central Laos, which represents the first recorded female we provide a redescription of C. bansocensis including expanded diagnosis. In addition, an updated identification key for the Cyrtodactylus known from Laos is provided.
... exhibited clear genetic and phenotypic differences from their sister lineages, H. jinpingensis (located more than 200 km away) and H. huishuiensis (located more than 350 km away), respectively, thus supporting the assumption of high local endemism for each karst region, even though some areas may have multiple sympatric species. High endemism in karst geckos is not only reported for Hemiphyllodactylus, but also for Cyrtodactylus (Davis et al., 2019;Grismer et al., 2021;Luu et al., 2016;Nazarov et al., 2018;Nguyen et al., 2017;Pauwels et al., 2016) and Cnemaspis (Grismer et al., 2014;Wood et al., 2017). In addition, high numbers of endemic flora and invertebrates have also been reported from limestone forest habitats (Clements et al., 2006;Marzuki et al., 2021;Nguyen et al., 2021). ...
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Karst habitats are hotspots of diversity and endemism. Their naturally fragmented distributions across broad geographic landscapes have led to the complex array of smaller evolutionary ecosystems that present unique challenges from a conservation perspective. Comprehensive biodiversity assessments of karst habitats have revealed that these ecosystems contain an almost unparalleled level of endemism, and many site-restricted species remain undescribed, thus posing considerable challenges for effective conservation management. Small rock-dwelling species, such as geckos, may be particularly prone to such isolation. In this paper, we discuss one such genus, i.e., Hemiphyllodactylus, and explore its diversity across karst landforms in Yunnan Province, southwestern China. Based on morphological and genetic data, we describe two new species of Hemiphyllodactylus from karst habitats in Simao District and Yanshan County. A phylogenetic tree for Hemiphyllodactylus was constructed using 1 039 base pairs (bp) of the mitochondrial NADH dehydrogenase subunit 2 gene ( ND2). The Simao and Yanshan specimens can be distinguished from all other congeners within their respective subclades based on uncorrected genetic pairwise distances greater than 6.3% and 4.3% respectively, as well as significant morphological differences. The discovery and description of these two new species brings the total number of described Hemiphyllodactylus species in China to 14 and indicates many more undescribed species from unsurveyed karst regions await discovery. Our findings suggest that karst ecosystems in Yunnan support a higher diversity of Hemiphyllodactylus than previously known. This study also highlights the importance of karst ecosystems as refugia for site-specific endemic species and the need for heightened conservation efforts.
... The following terms were abbreviated as shown in parentheses: snout-vent length (SVL), tail length (TL), tail width (TW), forearm length (FL), tibia length (TBL), axilla-togroin length (AG), head length (HL), head width (HW), head depth (HD), eye diameter (ED), eye to snout distance (ES), eyeto-nostril distance (EN), interorbital distance (IO), ear diameter (EL), internarial distance (IN), supralabials (SL), infralabials (IL), ventral scales (VS), subdigital lamellae on the fourth toe (ET4), femoral pores (FP), precloacal pores (PP), postprecloacal scales rows (PPS), body bands (BB), light bands on the tail (LB), and dark bands on the tail (DB). Morphological comparisons and analyses were based on specimen examination and data obtained from the existing literature (Bauer et al., 2009;Bauer et al., 2010;Kunya et al., 2014;Le et al., 2016;Liu and Rao, 2021;Luu et al., 2011;Nguyen et al., 2014;Nguyen et al., 2017;Nguyen et al., 2010;Pauwels et al., 2014a;Quang et al., 2007;Schneider et al., 2014;Sumontha et al., 2010;Tri, 2011;Zhang et al., 2021). ...
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Here, a new species of bent-toed gecko, Cyrtodactylus phukhaensis sp. nov., is described from Doi Phu Kha, Nan province, Thailand based on molecular and morphological evidence. A phylogeny based on NADH dehydrogenase subunit 2 (ND2) and its flanking tRNAs places the new species in the chauquangensis group as a sister taxon to Cyrtodact ylus wayakonei. The new species can be differentiated from other members of the chauquangensis group by having a unique combination of 7 or 8 supralabials, 23–28 longitudinal rows of dorsal tubercles, 8–10 infralabials, 9 femoral pores, 7 precloacal pores, 40–47 ventral scales, and a lack of bands crossing the temporal area. In addition, results from a chromosome study of C. phukhaensis sp. nov. showed that the new species has a diploid chromosome number of 40 with a fundamental number of 46. Te formula of the karyotype was as follows: 2n (40) = 2m + 4sm + 34t. Our findings suggest that further studies of Cyrtodactylus biodiversity in northern Tailand are needed.
... Bilateral scale counts were given as left/right. The methodology of measurements and meristic counts followed Liu and Rao (2021a, b): Morphological comparisons were based on the original descriptions of each species (Hoang et al. 2007;Bauer et al. 2009Bauer et al. , 2010Ngo and Grismer 2010;Nguyen et al. 2010Nguyen et al. , 2015bNguyen et al. , 2017Sumontha et al. 2010;Luu et al. 2011;Ngo 2011;Ngo and Chan 2011;Kunya et al. 2014;Nazarov et al. 2014;Nguyen et al. 2014;Schneider et al. 2014Schneider et al. , 2020Le 2016;Pham et al. 2019;Liu and Rao 2021a;Liu et al. 2021;Zhang et al. 2021). ...
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A new species of the Cyrtodactylus chauquangensis species group is described based on four specimens collected from the karst formations of Menglian County, Puer City, Yunnan Province, China. The new species can be separated from all other congeners by having a unique combination of morphological characters: a medium-sized body; ventrolateral folds present with interspersed small tubercles; seven precloacal pores in a continuous series in males, absent in females; enlarged femoral scales and femoral pores absent; two postcloacal tubercles on each side; and one or two rows of enlarged subcaudals. Genetically, the new species most closely related to C. wayakonei and the uncorrected sequence divergences of the ND2 gene and its flanking tRNAs between the new species and investigated congeners range from 7.2% to 18.4%.
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Cyrtodactylus is the most diverse genus of the family Gekkonidae and the world's third largest vertebrate ge-nus. The number of species has increased more than fourfold over the last two decades. Indochina, especially Vietnam and Laos, has witnessed a surge in new species discoveries over the last three decades. The species number reported from Laos and Vietnam has remarkably increased from five in 1997 to 71 species in 2021. However, within the genus, several taxonomic issues have not yet been fully resolved. Based on recently collected samples from Laos and Vietnam, we conducted a comprehensive molecular review of Cyrtodactylus A peer-reviewed open-access journal Hanh Thi Ngo et al. / ZooKeys 1097: 135-152 (2022) 136 occurring in Laos and Vietnam. Our molecular analysis with support from morphological comparisons showed that C. thuongae is a junior synonym of C. dati and C. rufford is a junior synonym of C. lomyenensis. In total, 68 described species distributed in Laos and Vietnam are undisputed with strong support from both molecular and morphological evidence. On the other hand, the molecular analyses revealed that there are at least seven undescribed species in Vietnam and Laos, one in the C. angularis group, one in the C. chauquangensis, and five in the C. irregularis group. This number will likely increase significantly, as previous work suggested that the C. angularis and C. irregularis groups harbor three and six unnamed lineages, respectively. Based on survey gaps identified in our study, it is clear that additional new species will be discovered in poorly studied regions of central Vietnam and northern and southern Laos. As many species in the genus are facing high extinction risks, several undescribed populations might already be severely threatened by human activities in both countries. Therefore, urgent taxonomic research is needed before conservation assessments of newly discovered taxa can be undertaken to protect them from anthropogenic threats.
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We describe a new species of Cyrtodactylus on the basis of two specimens collected from Ta Kou Nature Reserve, Binh Thuan Province, southern Vietnam. Cyrtodactylus chungi sp. nov. is distinguished from the remaining Indochinese bent-toed geckos by a combination of the following characters: relatively small body size (SVL up to 68.5 mm); a continuous neckband; 5 or 6 irregular transverse dorsal bands; 11 or 12 bands on original tail; keeled tubercles present on dorsum, posterior limbs and tail; 17 or 18 irregular dorsal tubercle rows; 30 or 31 ventral scale rows; ventrolateral skin folds indistinct; an angular series of seven precloacal pores in male and six pitted, enlarged precloacal scales in female, each series separated by a diastema of undifferentiated scales from 4-6 enlarged, poreless femoral scales; median subcaudals slightly enlarged; 17-20 subdigital lamellae under the fourth toe. Based on molecular analyses of the fragment of mitochondrial gene cytochrome c oxidase subunit I (COI), the new species is recovered as the sister taxon to Cyrtodactylus cattienensis s. str. with a genetic divergence of more than 9%. In phylogenetic analyses, the new species is recovered as a member of the Cyrtodactylus irregularis species group.
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