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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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27
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).
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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).
