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80
Accepted by A. Bauer: 29 May 2014; published: 9 Jul. 2014
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2014 Magnolia Press
Zootaxa 3835 (1): 080
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Article
http://dx.doi.org/10.11646/zootaxa.3835.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:3C8B1815-2A37-414C-99A9-982E274769AF
A new species of Cyrtodactylus (Squamata: Gekkonidae) from the karst forest of
northern Laos
NICOLE SCHNEIDER
1
, TRUONG QUANG NGUYEN
2,3,8
, MINH DUC LE
4,5,6
,
LIPHONE NOPHASEUD
7
, MICHAEL BONKOWSKI
2
& THOMAS ZIEGLER
1,2
1
AG Zoologischer Garten Köln, Riehler Straße 173, D-50735 Köln, Germany. E-mail: schneider.nicole87@googlemail.com and
ziegler@koelnerzoo.de
2
Zoological Institute, Department of Terrestrial Ecology, University of Cologne, Zülpicher Strasse 47b, D-50674 Cologne, Germany.
E-mail: m.bonkowski@uni-koeln.de
3
Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam.
Email: nqt2@yahoo.com
4
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
5
Centre for Natural Resources and Environmental Studies, Hanoi National University, 19 Le Thanh Tong, Hanoi, Vietnam
6
Department of Herpetology, American Museum of Natural History, Central Park West at 79
th
Street, New York, New York 10024
7
Faculty of Science, National University of Laos, Dong Dok Campus, Vientiane, Lao PDR. E-mail: nophasead2007@yahoo.com
8
Corresponding author
Abstract
We describe a new species of the gekkonid genus Cyrtodactylus on the basis of two specimens collected from limestone
forests of Luang Prabang Province, northern Laos. Morphologically, the new species is distinguishable from its congeners
by a combination of the following diagnostic characters: maximum SVL 86.1 mm; supralabials 9 or 10; infralabials 7–9;
dorsal tubercles in 15 or 16 rows at midbody; ventral scale rows 34–36 at midbody; precloacal groove absent; femoral
scales not distinctly enlarged; precloacal pores absent in females (unknown in males); subdigital lamellae under the fourth
finger 18 or 19, under the fourth toe 18–20; subcaudals not transversally enlarged; dorsal bands white, 4 or 5 between limb
insertions plus another one between hind limbs; tail banded. Based on molecular analyses, the new species is clustered in
the same clade with C. wayakonei and two other species from Luang Prabang and Houaphan provinces.
Key words: Bent-toed gecko, limestone forest, phylogeny, taxonomy, Luang Prabang Province
Introduction
With 186 recognized species, Cyrtodactylus is one of the most diverse genera of lizards in terms of species richness
(Uetz & Hosek 2014). Since 2010, 63 species have been described in the genus and most of these new discoveries
were from Southeast Asian countries (Nguyen et al. 2013; Ziegler et al. 2013; Grismer et al. 2014; Luu et al. 2014;
Phung et al. 2014; Pauwels & Sumontha 2014; Pauwels et al. 2013, 2014; Schneider et al. 2014; Uetz & Hosek
2014). A major factor for the high species diversity of bent-toed geckos in Southeast Asia is the presence of a large
area of tropical rain forest and a number of species complexes. For instance, 10 new species have been described
within the Cyrtodactylus irregularis species complex from the Indochina region and eight new species were
discovered in the C. pulchellus species complex from the Thai-Malay Peninsula (Grismer et al. 2012; Sumontha et
al. 2012; Nguyen et al. 2013; Schneider et al. 2014). In Laos, nine species of Cyrtodactylus are currently
recognized and seven of them have been recorded from limestone karst forests, namely C. interdigitalis Ulber,
1993 and C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, 2011 from Vientiane Province; C.
lomyenensis Ngo & Pauwels, 2010, C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010,
and C. jaegeri Luu, Calame, Bonkowski, Nguyen & Ziegler, 2014 from Khammuane Province; C. wayakonei
Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 from Luang Nam Tha Province; and C. teyniei David, Nguyen,
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NEW CYRTODACTYLUS FROM LAOS
Schneider & Ziegler, 2011 from Borikhamxay Province (Teynié & David 2010; Luu et al. 2013). During recent
field work in northern Laos, we collected two specimens of an unnamed species of Cyrtodactylus in the karst forest
of Luang Prabang Province. Based on morphological examination and molecular phylogenetic data, we herein
describe it as a new species from northern Laos.
Material and methods
Sampling. Field surveys were conducted in August 2013 in Luang Prabang District, Luang Prabang Province, Lao
People’s Democratic Republic. Tissue samples were preserved separately in 95% ethanol and voucher specimens
were fixed in approximately 85% ethanol, then later transferred to 70% ethanol for permanent storage. Specimens
were subsequently deposited in the collections of the Institute of Ecology and Biological Resources (IEBR), Hanoi,
Vietnam and the National University of Laos (NUOL), Vientiane, Laos. Other abbreviations are as follows: MHNG
= Muséum d'Histoire Naturelle, Genève, Switzerland; MTD = Senckenberg Naturhistorische Sammlungen
Dresden, Museum für Tierkunde, Dresden, Germany; NEM = Namlik Ecovillage Museum, Ban That Wang Monh,
Vientiane Province, Laos; VNUH = Zoological Museum, University of Natural Science, Vietnam National
University, Hanoi, Vietnam; and ZFMK = Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany.
Molecular data and phylogenetic analyses. We included newly collected samples of nine Cyrtodactylus
species from Laos (Table 1) and of 13 species from Vietnam. A member of the C. pulchellus Gray, 1828 complex
was used as an outgroup.
TABLE 1. Additional samples from Laos used in molecular analyses (for abbreviations see Material and Methods).
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). Data were analysed using maximum parsimony (MP) and Bayesian analyses, following the
same procedures as described in Schneider et al. (2014). The optimal model for nucleotide evolution was set to
GTR+I+G as selected by Modeltest v3.7 (Posada & Crandall 1998). The cut off point for the burn-in function was
set to 25 in the Bayesian analysis, as -lnL scores reached stationarity after 25,000 generations in both runs. Nodal
support was evaluated using Bootstrap replication (BP) as calculated in PAUP and posterior probability (PP) in
MrBayes v3.2. Uncorrected pairwise divergences were calculated in PAUP*4.0b10 (Table 2).
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
(TL), 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; eye to ear distance (EE), from
posterior corner of eye to anterior margin of ear opening; internarial distance (IND), measured between inner
border 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
Species GenBank number Locality Voucher information
Cyrtodactylus sp1 KJ817428 Houphan Province IEBR A.2013.109
Cyrtodactylus sp2 KJ817432 Luang Prabang Province IEBR A.2013.110
Cyrtodactylus sp2 KJ817433 Luang Prabang Province IEBR A.2013.111
C. lomyenensis KJ817436 Khammuane Province IEBR KM2012.54
C. pageli KJ817431 Vientiane Province ZFMK 91827
C. sp3 KJ817429 Khammuane Province IEBR A.2013.89
C. sp4 KJ817437 Khammuane Province IEBR A.2013.112
C. teyniei KJ817430 Khammuane Province IEBR KM2012.77
C. vilaphongi sp. nov. KJ817434-5 Luang Prabang Province NUOL R-2013.5, IEBR A.2013.103
C. wayakonei KJ817438 Luang Nam Tha Province ZFMK 91016
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elbow; shank length (SL), from the base of heel to the knee; length of first finger (LeF1) and of fourth finger
(LeF4), from the base to the tip of finger; length of first toe (LeT1) and of fourth toe (LeT4), from the base to the
tip of toe; rostral width (RW); rostral heigth (RH); mental width (MW); mental length (ML). Scale counts were
taken as follows: supralabials (SPL); infralabials (IFL); nasal scales surrounding nare, from rostral to labial
(excluding rostral and labial), i.e. nasorostral, supranasal, postnasals (N); postrostrals or internasals (I); scale rows
between fifth supralabials (SC5SPL); interorbitals (IO), scales between the anterior corners of the eyes; spinous
ciliaries (CS); postmentals, i.e. scales bordering mental shield, except infralabials (PM); gulars bordering the
postmentals (GP); granular scales surrounding dorsal midbody tubercles (GSDT); dorsal tubercle rows at midbody
(DTR); ventral scales in longitudinal rows at midbody (V); number of scales along the midbody from mental shield
to anterior edge of cloaca (SMC); scales around midbody (MS); enlarged femoral scales (EFS); precloacal pores
(PP); femoral pores (FP); total number of postcloacal tubercles (PAT); subdigital lamellae under the first finger
(LF1); subdigital lamellae under the fourth finger (LF4); subdigital lamellae under the first toe (LT1); subdigital
lamellae under the fourth toe (LT4). Bilateral scale counts were given as left/right.
Results
Phylogenetic analyses. We successfully sequenced 658 bps for 11 newly collected samples. The combined matrix
contained 658 aligned characters with no gap. Missing segments in shorter sequences were specified as missing
data in the matrix. MP analysis of the dataset recovered six most parsimonious trees with 1606 steps (CI = 0.32; RI
= 0.60). The topology within the large clade comprising a majority of species from Vietnam was similar to the one
generated in a previous study (Schneider et al. 2014) (Fig. 1). Species from Laos are grouped in two independent
clades with strong support from both MP and Bayesian analyses for one clade (BP = 94, PP = 100), and strong
support from the Bayesian analysis only for another (BP = 57, PP = 99) (Fig 1). The new species is strongly
corroborated as a sister taxon to Cyrtodactylus sp1 from Houaphan Province (BP = 100, PP = 100). It is also
significantly divergent from others in terms of genetic distance with a minimum pairwise divergence of
approximately 8% in the mitochondrial fragment of COI (Table 2).
TABLE 2. Uncorrected (“p”) distance matrix showing percentage pairwise genetic divergence (COI) between Laotian
Cyrtodactylus species. Note: The minimal genetic divergence between C. vilaphongi sp. nov. and species within the
clade containing mostly species from Vietnam is approximately 17%. Details of the divergence within this clade can be
found in Schneider et al. (2014).
Morphological comparisons. Although only two female specimens were available for morphological
comparisons, and consequently some characters that are usually present in males, are still unknown (e.g., femoral
and precloacal pores), the undescribed species from Laos is clearly differentiated from the remaining congeners of
the genus Cyrtodactylus by a unique suite of features. We compare the unnamed species from Laos with all other
members of Cyrtodactylus from southern China and mainland Southeast Asia based on examination of specimens
(see Appendix) and data obtained from the literature.
Species 123456789
1. Cyrtodactylus sp1 (KJ817428) -
2. Cyrtodactylus sp2 (KJ817432-3) 13.1 -
3. C. lomyenensis (KJ817436) 20.8 20.1 -
4. C. pageli (KJ817431) 19.3 20.4 17.3 -
5. C. sp3 (KJ817429) 21.9 20.4 14.4 18.9 -
6. C. sp4 (KJ817437) 19.5 18.7 17.9 16.4 17.0 -
7. C. teyniei (KJ817430) 20.8 21.0 14.6 17.5 14.4 17.8 -
8. C. vilaphongi sp. nov. (KJ817434-5) 8.21 12.9 21.0 19.3 20.5 20.1 18.1 -
9. C. wayakonei (KJ817438) 17.0 15.2 20.7 20.4 19.6 20.4 21.1 15.8 -
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FIGURE 1. One of the six most parsimonious maximum parsimony trees based on the partial COI gene. Numbers above and
below branches are bootstrap values (>50%) and Bayesian posterior probabilities, respectively. The bracket and number
indicate the sister relationship between two taxa and the posterior probability supported by the Bayesian analysis, respectively.
In the Bayesian tree, C. taynguyenensis + C. pseudoquadrivirgatus + C. kingsadai + C. cryptus are grouped with C. cattienensis
+ C. phuocbinhensis with weak support (PP = 60%). LP: Luang Prabang.
The new species can be distinguished from the following species by the absence of transversely enlarged
subcaudals: C. angularis (Smith, 1921), C. annandalei Bauer, 2003, C. auribalteatus Sumontha, Panitvong &
Deein, 2010, C. ayeyarwadyensis Bauer, 2003, C. badenensis Nguyen, Orlov & Darevsky, 2006, C. bichnganae
Ngo & Grismer, 2010, C. brevipalmatus (Smith, 1923), C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva &
Nguyen, 2007, C. chanhomeae Bauer, Sumontha & Pauwels, 2003, C. chauquangensis Hoang, Orlov, Ananjeva,
Johns, Hoang & Dau, 2007, C. condorensis (Smith, 1920), C. consobrinus (Peters, 1871), C. cucphuongensis Ngo
& Chan, 2011, C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, 2010, C.
eisenmanae Ngo, 2008, C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome &
Kunya, 2009, C. feae (Boulenger, 1893), C. grismeri Ngo, 2008, C. hontreensis Ngo, Grismer & Grismer, 2008, C.
huongsonensis Luu, Nguyen, Do & Ziegler, 2011, C. interdigitalis, C. intermedius (Smith, 1917), C. jaegeri, C.
jarujini Ulber, 1993, C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda, 2014, C. kingsadai
Ziegler, Phung, Le & Nguyen, 2013, C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer,
Wangkulangkul, Grismer & Pauwels 2012, C. lomyenensis, C. martini Ngo, 2011, C. nigriocularis Nguyen, Orlov
& Darevsky, 2006, C. oldhami (Theobald, 1876), C. pageli, C. paradoxus (Darevsky & Szczerbak, 1997), C.
peguensis (Boulenger, 1893), C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, 2002, C. phuketensis
Sumontha, Pauwels, Kunya, Nitikul, Samphanthamit & Grismer, 2012, C. phuquocensis Ngo, Grismer & Grismer,
2010, C. pulchellus Gray, 1828, C. roesleri, C. rubidus (Blyth, 1861), C. russelli Bauer, 2003, C. samroiyot
Pauwels & Sumontha, 2014, C. sanook Pauwels, Sumontha, Latinne & Grismer, 2013, C. slowinskii Bauer, 2002,
C. sumonthai Bauer, Pauwels & Chanhome, 2002, C. surin Chan-ard & Makchai, 2011, C. takouensis Ngo &
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Bauer, 2008, C. teyniei, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome 2004, C. thochuensis Ngo &
Grismer, 2012, C. tigroides Bauer, Sumontha & Pauwels, 2003, C. variegatus (Blyth, 1859), and C. yangbayensis
Ngo & Chan, 2010.
The specimens of Cyrtodactylus from Laos differ from C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung,
Nguyen, Hoang & Ziegler, 2012 by having fewer ventral scale rows (34–36 vs. 38–43), fewer dorsal tubercle rows
(15 or 16 vs. 18–24), the absence of enlarged femoral scales and enlarged tubercles on lateral fold (present in C.
bidoupimontis), and the presence of light transversal dorsal bands (dark dorsal bands in C. bidoupimontis).
The new species is distinguishable from C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang
& Ziegler, 2012 and C. irregularis (Smith, 1921) by having fewer ventral scales (34–36 vs. 36–46 and 38–45,
respectively), more subdigital lamellae under the fourth finger (18 or 19 vs. 15–17 and 15 or 16, respectively), the
presence of light dorsal bands (absent in C. bugiamapensis and C. irregularis), and the absence of enlarged femoral
scales (present in C. bugiamapensis and C. irregularis).
The new species differs from C. buchardi David, Teynié & Ohler, 2004 by having fewer dorsal tubercle rows
(15 or 16 vs. 25), more ventral scales (34–36 versus 30), and more subdigital lamellae under the fourth finger and
toe (18 or 19 and 18–20 vs. 14 and 12, respectively).
The new species differs from C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler,
2009, C. dati Ngo, 2013 and C. guakanthanensis Grismer, Belabut, Quah, Chan, Wood & Hasim, 2014 by the
absence of distinctly enlarged femoral scales. In addition, the new species has more subdigital lamellae under the
fourth finger than C. cattienensis (18 or 19 vs. 12–16) and has fewer ventral scales than C. dati and C.
guakanthanensis (34–36 vs. 42–48 and 37–44, respectively).
The new species has 34–36 ventral scales, thus it differs from the following species: C. aequalis Bauer, 2003
(24), C. brevidactylus Bauer, 2002 (45), C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007 (47–50), C.
phuocbinhensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen, Nguyen, Hoang, Che, Murphy &
Zhang, 2013 (43–47), and C. taynguyenensis Nguyen, Le, Tran, Orlov, Lathrop, Macculloch, Le, Jin, Nguyen,
Nguyen, Hoang, Che, Murphy & Zhang, 2013 (42–49). In addition, the new species differs from C. phuocbinhensis
by the absence of enlarged femoral scales (present in C. phuocbinhensis) and by having a different dorsal color
pattern (dark dorsum with light bands vs. dark stripes and blotches on brighter dorsum) and from C. taynguyenensis
by having a different dorsal color pattern (light bands vs. dark blotched or irregular bands on dorsum). The new
species further differs from C. cryptus by having fewer subdigital lamellae under the fourth toe (18–20 vs. 20–23)
and a different dorsal color pattern (light narrow bands vs. dark broad bands).
The new species is distinguishable from C. huynhi Ngo & Bauer, 2008 by having fewer ventral scales (34–36
vs. 43–46), more subdigital lamellae under the fourth finger (18 or 19 vs. 14–17), and the absence of enlarged
femoral scales (present in C. huynhi).
The new species is distinguishable from C. thuongae Phung, van Schingen, Ziegler & Nguyen, 2014 by the
absence of enlarged femoral scales (vs. present) and by having a different dorsal color pattern (banded vs.
blotched).
The new species is distinguishable from C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008 by having fewer
dorsal tubercle rows (15 or 16 vs. 20–24) and the presence of light dorsal bands (dark dorsal bands in C. ziegleri).
The new species has 15 or 16 dorsal tubercle rows, thus it differs from C. gansi Bauer, 2003 (20–25), C.
tamaiensis Smith, 1940 (21), C. wakeorum Bauer 2003 (24), and C. wayakonei (17–19).
The new species has no precloacal pores in females, thus it differs from the following species, which have
precloacal pores or pitted precloacal scales in females: C. chrysopylos Bauer, 2003 and C. pseudoquadrivirgatus
Rösler, Vu, Nguyen, Ngo & Ziegler, 2008 and C. quadrivirgatus Taylor, 1962. The new species further differs from
C. quadrivirgatus by having fewer ventral scales (34–36 vs. 40) and the absence of enlarged femoral scales.
Cyrtodactylus vilaphongi sp. nov.
(Figs. 2–4)
Holotype. IEBR A.2013.103, adult female, collected by Truong Quang Nguyen and Vilaphong Kanyasone on 16
August 2013, near Ban Xieng Muak (19
o
48.772’N, 102
o
06.181’E, elevation 597 m), Luang Prabang District,
Luang Prabang Province, Laos.
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FIGURE 2. Dorso-lateral view of (A) the holotype of Cyrtodactylus vilaphongi sp. nov. (IEBR A.2013.103) and (B) the
paratype (NUOL R-2013.5) in life from Luang Prabang Province, Laos. Photos by T. Q. Nguyen.
Paratype. NUOL R-2013.5, adult female, collected by Truong Quang Nguyen, Nicole Schneider, Liphone
Nophaseud, Vilaphong Kanyasone on 10 August 2013, the same locality as the holotype.
Diagnosis. The new species can be distinguished from all congeners on the basis of the following combination
of characters: maximum SVL 86.1 mm, supralabials 9 or 10; infralabials 7–9; dorsal tubercles in 15 or 16 rows at
midbody; ventral scale rows 34–36 at midbody; precloacal groove absent; femoral scales not distinctly enlarged;
precloacal pores absent in females (unknown in males); subdigital lamellae under the fourth finger 18 or 19, under
the fourth toe 18–20; subcaudals not transversally enlarged; dorsal bands white, 4 or 5 between limb insertions plus
another one between hind limbs; tail banded.
Description of the holotype. Adult female, SVL 86.1 mm, tail regenerated (TL 61.2 mm); rostral square-
shaped, wider than high (RW 4.1 mm, RH 2.4 mm), medially with a straight, vertical suture, in contact with
nasorostral, nare, and first supralabial on each side; supralabials 9 or 10; supralabials separated from orbit by 3 or 4
rows of scales; nares in contact with rostral, nasorostral, supranasal, two postnasals, and first supralabial;
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internasals absent; scales on snout distinctly larger than on upper head; loreal region slightly upraised; scales
between fifth supralabials across the dorsal head surface in 30 rows; scales between anterior corners of eyes 32;
interorbital region with small, round, convex scales, outer ones more oval; scales in postorbital region distinctly
smaller (ca. half the size) than snout scales, round and convex, interrupted by a few small tubercles, in 3 or 4 rows
laterally; dorsal surface of head without enlarged tubercles; pupil vertical; spinous ciliaria absent; ear opening
vertical, oval; mental triangular, slightly narrower than rostral (MW: 3.6 mm), in contact with two postmentals and
the first infralabial on each side; infralabials 8 or 9; postmentals surrounded by first infralabial on each side and six
granular scales posteriorly, two outer ones enlarged; gular scales granular.
FIGURE 3. Dorsal view (A), ventral view (B), and lateral view (C) of the head of the holotype of Cyrtodactylus vilaphongi sp.
nov. (IEBR A.2013.103) in preservative. Photos N. Schneider.
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FIGURE 4. Precloacal region and ventral tail (A) and dorsal scales at midbody (B) of the holotype of Cyrtodactylus vilaphongi
sp. nov. (IEBR A.2013.103) in preservative. Photos N. Schneider.
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Dorsal scales small; dorsal tubercles in 15 longitudinal rows at midbody, separated from each other by 2 or 3
dorsal scales and surrounded by 9–11 granular scales; tubercles on flanks slightly smaller than dorsal tubercles;
ventrolateral folds slightly developed, with tubercles; ventral scales round, imbricated, 2 or 3 times larger than
gular and throat scales, twice the size of dorsal scales; ventral scales 34, plus 122 dorsal scales around midbody;
scales between mental and cloacal slit 165; dorsal surface of limbs with tubercles, distinctly higher counts on hind
limbs; distinctly enlarged femoral scales absent; fingers and toes free of webbing; relative finger lengths
I<V<II<IV<III, relative toe lengths I<II<V<IV<III; claw bordered by two scales; subdigital lamellae: finger I 15
(with 3 basally broadened lamellae), finger II 14–16 (4), finger III 18 (5), finger IV 18 or 19 (5), finger V 18 (5),
toe I 14 or 15 (3), toe II 17 or 18 (4 or 5), toe III 19 (5), toe IV 18 or 19 (5 or 6), toe V 20 or 21 (5 or 6); precloacal
depression absent; precloacal pores absent; enlarged precloacal scales 5, in one row; adjoining scales in precloacal
region twice the size of femoral scales or ventrals; postcloacal tubercles 2 on each side, well developed; dorsal tail
base with tubercles, regenerated part without tubercles or whorls; subcaudals of original part of tail not
transversally enlarged.
Coloration in life: Dorsal head dark brown, marbled in light yellow; each side with a blackish brown stripe
stretching from the posterior corner of the eye to the neck, connecting with the neck band on the right side,
interrupted on the left side; irregular dark blotches with light margins present between eyes, on occiput and on
postorbital region; ciliaria bright yellow; iris yellowish brown, pupil slit edged in orange; neck band blackish
brown, widened posteriorly, edged in yellow anteriorly and posteriorly; ground color of dorsum blackish brown
with five transverse yellowish white bands between limb insertions; flanks and upper limbs with yellowish white
spots, in irregular rows; tail base dark brown with a white band, regenerated part uniformly dark brown; ventral
surface light beige.
Va ri at io n . The paratype largely corresponds with the description of the holotype. For measurements,
scalation, and color pattern variation see Fig. 2 and Table 3. Neck band is not interrupted in the paratype. The
original tail has nine white dorsal bands plus a bright tip, dark brown ventrally, with dorsal tubercles at the base
only, and subcaudals not transversally enlarged.
TABLE 3. Morphological characters of Cyrtodactylus vilaphongi sp. nov. from Luang Prabang Province, Laos (* =
regenerated tail, other abbreviations defined in the text).
Characters IEBR A.2013.103 (female) NUOL R-2013.5 (female)
Measurements (mm)
SVL 86.1 60.9
TL 61.2* 68.1
AG 36.6 25.1
HL 23.6 16.6
HW 17.4 13.0
HH 9.7 6.9
SE 10.2 7.5
EE 7.4 4.9
IND 2.9 2.5
IOD 6.7 5.6
OD 5.5 4.1
ED 1.7 1.6
ForeaL 13.2 10.5
SL 14.9 12.4
LeF4 7.5 5.5
LeT4 9.6 6.8
RW 4.1 2.9
......continued on the next page
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Distribution. Cyrtodactylus vilaphongi is currently known only from the type locality in Luang Prabang
Province, Laos (Fig. 5).
Etymology. The new species is named in honour of Mr. Vilaphong Kanyasone, a staff of Provincial Natural
Resources and Environmental Office of Luang Prabang Province, Laos, who collected the holotype and provided a
great support for our field research in Laos.
Ecological notes. Cyrtodactylus vilaphongi inhabits disturbed secondary limestone forest near a residential
area at elevations between 577 and 597 m a.s.l. Specimens were found at night, on karst cliff and rock boulder ca.
0.5 m above the ground (Fig. 6). Air temperature at the site was 24–30
o
C and humidity ranged from 72 to 86%.
Discussion
Morphologically, Cyrtodactylus vilaphongi is most similar to C. wayakonei from Luang Nam Tha Province,
northern Laos in its size and scalation. Both species are limestone-dwelling bent-toed geckos and the distance
between the type localities of these species is approximately 130 km. However, C. vilaphongi can be distinguished
from C. wayakonei by having fewer dorsal tubercle rows (15 or 16 vs. 17–19), more scales around midbody
(106–122 vs. 85–98), and the presence of narrow yellowish white bands on its dorsum (vs. irregular purplish brown
bands or reticulated pattern) (Nguyen et al. 2010). Nonetheless, C. vilaphongi is distantly related to C. wayakonei
TABLE 3. (Continued)
Characters IEBR A.2013.103 (female) NUOL R-2013.5 (female)
RH 2.4 1.9
MW 3.6 2.5
ML 2.5 2
Scalation
SPL 9/10 9/9
IFL 9/8 8/7
N33
I00
SC5SPL 30 45
IO 32 31
CS 0 0
PM 2 2
GP 6 6
DTR 15 16
GSDT 9–11 8–10
SMC 165 161
MS 122 106
V3436
LF1 13/15 12/13
LF4 18/19 18/19
LT1 14/15 13/12
LT4 19/18 18/20
PP 0 0
FP 0 0
PFP 0 0
PAT 2/2 2/2
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based on the molecular data, which suggests the adaptation to limestone habitat as morphological convergence
evolution.
FIGURE 5. Type locality (black dot) of Cyrtodactylus vilaphongi in Luang Prabang Province, Laos.
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TABLE 4. Morphological comparison of Cyrtodactylus species without enlarged subcaudals from Laos and neighboring countries (after Ziegler et al. 2013; Nguyen et al.
2013; Phung et al. 2014; Schneider et al. 2014; max. = maximum; – = data unavailable; other abbreviations defined in the text).
SVL TaL PP PP
Species (mm) (mm) V EFS FP
(males) (females) LD4 LT4 Color pattern of
dorsum
Cyrtodactylus vilaphongi sp. nov. 60.9–86.1 61.2–68.1 34–36 0 – – 0 18–19 18–20 banded
C. bidoupimontis 74.0–86.3 75.0–86.0 38–43 6–8 absent 4–6 0 15–20 18–23 banded
C. bugiamapensis 58.6–76.8 65.3–83.0 36–46 6–10 absent 7–8 0–7 15–17 17–20 blotched
C. buchardi 60.0–65.0 46.0–54.0 30 absent absent 9 0 14 12 blotched
C. cattienensis 43.5– 69.0 51.0–64.7 28–42 3–8 absent 6–8 6–7 12–16 14–19 banded
C. cryptus 62.5–90.8 63.5–88.4 47–50 absent absent 9–11 9–11 18–19 20–23 banded
C. cucdongensis 55.8–65.9 max. 81.3 40–44 5–9 absent 5–6 4–6 13–18 15–20 banded
C. huynhi 54.8–79.8 61.5–78.6 43–46 3–5 3–8 7–9 0–8 14–17 17–21 banded
C. irregularis 72.0–86.0 66.0–74.0 38–45 7–8 – 5–7 0–6 15–16 18–19 blotched
C. phuocbinhensis 46.0–60.4 76.1 43–47 5 absent 7 0 16–21 17–19 striped/blotched
C. pseudoquadrivirgatus 48.6–83.3 55.7–82.3 41–57 absent absent 5–9 5–10 15–21 16–25 blotched
C. quadrivirgatus 39.0–67.0 77 40 present absent 4 4 – – striped
C. taynguyenensis 60.0–85.0 66.0–94.0 42–49 absent absent 6 0 13–18 17–21 blotched/banded
C. thuongae 57.3–77.6 max. 78.1 29–44 2–5 0–3 0–1 0 14–17 14–20 blotched
C. wayakonei 72.0–86.8 76.8–89.0 31–35 absent absent 6–8 7 17–18 19–20 banded
C. ziegleri 84.6–93.0 95.0–107.0 33–39 8–10 0–6 5–8 0–8 16–19 18–21 banded
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FIGURE 6. General vegetation type of the karst forest (A) and biotope (B) of Cyrtodactylus vilaphongi in Luang Prabang
Province, Laos. Photos T. Q. Nguyen.
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Acknowledgements
We are grateful to S. Wayakone, S. Bounphanmy, B. Praxaysombath (Vientiane) and V. Kanyasone (Luang
Prabang) for supporting our field research in Laos. Export of collected specimens was done due to the export
permit Nr. 139/13 signed by the CITES Management Authority of Lao PDR. T. Q. Nguyen thanks A. Teynié
(Aubière), C. X. Le, T. H. Tran (Hanoi), T. Pagel (Cologne), W. Böhme and D. Rödder (Bonn) for support of his
research. Thanks to H. T. Duong (Hanoi) for laboratory assistance. We are grateful to A. Bauer (Villanova), S. N.
Nguyen (Ho Chi Minh City), and O. S. G. Pauwels (Brussels) for reviewing the manuscript. Thanks to E. Sterling
(New York) and K. Koy (Berkeley) for providing the map. Field survey in Luang Prabang Province was funded by
the Alexander von Humboldt Foundation for TQN (VIE 1143441) and Cologne Zoo for NS. Research of T. Q.
Nguyen was funded by the Alexander von Humboldt Foundation (VIE 1143441).
References
Bauer, A.M. (2002) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from Myanmar. Proceedings of the California
Academy of Sciences, 53, 73–86.
Bauer, A.M. (2003) Descriptions of seven new Cyrtodactylus (Squamata: Gekkonidae) with a key to the species of Myanmar.
Proceedings of the California Academy of Sciences, 54, 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.M., 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.
Blyth, E. (1859) Report of the Curator, Zoological Department, for February to May meetings, 1859. Journal and Proceedings
of the Asiatic Society of Bengal, 28, 271–298.
Blyth, E. (1861) Proceedings of the Society. Report of the Curator. Journal and Proceedings of the Asiatic Society of Bengal,
xxix (1860), 87–115.
Boulenger, G.A. (1893) Concluding report on the reptiles and batrachians obtained in Burma by Signor L. Fea, dealing with a
collection made in Pegu and Karin Hills in 1887–1888. Annali del Museo civico di storia naturale di Genova, 2, 303–347.
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, 7–15.
Darevsky, I.S. & Szczerbak, N.N. (1997) A new gecko of the genus Gonydactylus (Sauria: Gekkonidae) with a key to the
species from Vietnam. Asiatic Herpetological Research, 7, 19–22.
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.
David, P., Teynié, A. & Ohler, A. (2004) A new species of Cyrtodactylus Gray 1827 (Reptilia: Squamata: Gekkonidae) from
southern Laos. The Raffles Bulletin of Zoology, 52, 621–627.
Geissler, P., Nazarov, R., Orlov, N.L., Böhme, W., Phung, T.M., Nguyen, T.Q. & Ziegler, T. (2009) A new species of the
Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa, 2161, 20–32.
Gray, J.E. (1828) A synopsis of the genera of Saurian reptiles in which some new genera are indicated, and the others reviewed
by actual examination. Philosophical Magazine, London, 2, 54–58.
Grismer, L.L., Belabut, D.M., Quah, E.S.H., Chan, K.O., Wood, P.L. & Hasim, R. (2014) A new species of karst forest-adapted
Bent-toed Gecko (genus Cyrtodactylus Gray, 1827) belonging to the C. swoderi complex from a threatened karst forest in
Perak, Peninsular Malaysia. Zootaxa, 3755 (5), 434–446.
http://dx.doi.org/10.11646/zootaxa.3755.5.3
Grismer, L.L., Wood, P.L., Quah, E.S.H., Anuar, S., Muin, M.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.
Heidrich, A., Rösler, H., Vu, N.T., Böhme, W. & Ziegler, T. (2007) Another new Cyrtodactylus (Squamata: Gekkonidae) from
Phong Nha – Ke Bang National Park, central Truong Son, Vietnam. Zootaxa, 1445, 35–48.
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
SCHNEIDER ET AL.
94
·
Zootaxa 3835 (1) © 2014 Magnolia Press
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.
Le, M., Raxworthy, C.J., McCord, W.P. & Mertz, L. (2006) A molecular phylogeny of tortoises (Testudines: Testudinidae)
based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 40, 517−531.
http://dx.doi.org/10.1016/j.ympev.2006.03.003
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.
http://dx.doi.org/10.11646/zootaxa.3760.1.3
Luu, V.Q., Nguyen, T.Q., Calame, T., Hoang, T.T., Southichack, S., Bonkowski, M. & Ziegler, T. (2013) New country records
of reptiles from Laos. Biodiversity Data Journal, 1, 1−14.
http://dx.doi.org/10.3897/bdj.1.e1015
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.
Mahony, S. (2009) Taxonomic status of Cyrtodactylus khasiensistamaiensis (Smith, 1940) and description of a new species
allied to C. chrysopylos Bauer, 2003 from Myanmar (Reptilia: Gekkonidae). Hamadryad, 34, 62–74.
Nazarov, R.A., Orlov, N.L., Nguyen, S.N. & Ho, C.T. (2008) Taxonomy of naked-toe geckos Cyrtodactylus irregularis
complex of South Vietnam and description of a new species from Chu Yang Sin Natural Park (Krong Bong District, Dac
Lac Province, Vietnam). Russian Journal of Herpetology, 15, 141–156.
Nazarov, R., Poyarkov, N.A., Orlov, N.L., Phung, T.M., Nguyen, T.T., Hoang, D.M. & Ziegler, T. (2012) Two new cryptic
species of the Cyrtodactylus irregularis complex (Squamata: Gekkonidae) from southern Vietnam. Zootaxa, 3302, 1–24.
Ngo, T.V. (2008) Two new cave-dwelling species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Southwestern Vietnam.
Zootaxa, 1909, 37–51.
Ngo, T.V. (2011) Cyrtodactylus martini, another new karst-dwelling Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from
Northwestern Vietnam. Zootaxa, 2834, 33–46.
Ngo, T.V. (2013) Cyrtodactylus dati, a new forest dwelling Bent-toed Gecko (Squamata: Gekkonidae) from southern Vietnam.
Zootaxa, 3616 (2), 151–164.
http://dx.doi.org/10.11646/zootaxa.3616.2.4
Ngo, T.V. & Bauer, A.M. (2008) Descriptions of two new species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae)
endemic to southern Vietnam. Zootaxa, 1715, 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, T.V. & Grismer, L.L. (2012) A new endemic species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu
Island, southwestern Vietnam. Zootaxa, 3228, 48–60.
Ngo, T.V., 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, T.V., 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
Khammuane Province, southern Laos. Zootaxa, 2730, 44–56.
Nguyen, S.N., Le, N.T.T., Tran, D.T.A., Orlov, N.L., Lathrop, A., Macculloch, R.D., Le, D.T.T., Jin, J.-Q., Nguyen, L.T.,
Nguyen, T.T., Hoang, D.D., Che, J., Murphy, R.W. & Zhang, Y.-P. (2013) Phylogeny of the Cyrtodactylus irregularis
species complex (Squamata: Gekkonidae) from Vietnam with the description of two new species. Zootaxa, 3737 (4),
399–414.
http://dx.doi.org/10.11646/zootaxa.3737.4.4
Nguyen, S.N., Orlov, N.L. & Darevsky, I.S. (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, T.Q., Kingsada, P., Rösler, H., Auer, M. & Ziegler, T. (2010) A new species of Cyrtodactylus (Squamata: Gekkonidae)
from northern Laos. Zootaxa, 2652, 1–16.
Orlov, N.L., Nguyen, Q.T., Nazarov, R.A., Ananjeva, 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].
Russian Journal of Herpetology, 14, 145–152.
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.
http://dx.doi.org/10.11646/zootaxa.3755.6.4
Zootaxa 3835 (1) © 2014 Magnolia Press
·
95
NEW CYRTODACTYLUS FROM LAOS
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.
http://dx.doi.org/10.11646/zootaxa.3635.3.7
Pauwels, O.S.G., Sumontha, M., Panitvong, N. & Varaguttanonda, V. (2014) Cyrtodactylus khelangensis, a new cave-dwelling
Bent-toed Gecko (Squamata:Gekkonidae) from Lampang Province, northern Thailand. Zootaxa, 3755 (6), 584–594.
http://dx.doi.org/10.11646/zootaxa.3755.6.5
Peters, W. (1871) Über neue Reptilien aus Ostafrika und Sarawak (Borneo), vorzüglich aus der Sammlung des Hrn. Marquis J.
Doria zu Genua. Königlich Preussische Akademie der Wissenschaften zu Berlin, 566−581.
Phung, T.M., van Schingen, M., Ziegler, T. & Nguyen, T.Q. (2014) A third new Cyrtodactylus (Squamata: Gekkonidae) from
Ba Den Mountain, Tay Ninh Province, southern Vietnam. Zootaxa, 3764 (3), 347−363.
http://dx.doi.org/10.11646/zootaxa.3764.3.5
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817−818.
http://dx.doi.org/10.1093/bioinformatics/14.9.817
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. &
Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology, 61, 539−542.
http://dx.doi.org/10.1093/sysbio/sys029
Rösler, H., Vu, T.N., Nguyen, T.Q., Ngo, T.V. & Ziegler, T. (2008) A new Cyrtodactylus (Squamata: Gekkonidae) from central
Vietnam . Hamadryad, 32, 125–140.
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.
Schneider, N., Phung, T.M., Le, M.D., Nguyen, T.Q. & Ziegler, T. (2014) A new Cyrtodactylus (Squamata: Gekkonidae) from
Khanh Hoa Province, southern Vietnam. Zootaxa, 3785 (4), 518–532.
http://dx.doi.org/10.11646/zootaxa.3785.4.2
Smith,M.A. (1917) Descriptions of new reptiles and a new batrachian from Siam. Journal of the Natural History Society of
Siam, 2, 221–225.
Smith, M.A. (1920) Reptiles and batrachians collected on Pulo Condore. Journal of Natural History Society of Siam, 4, 93–97.
Smith, M.A. (1921) 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. (1923) Notes on reptiles and batrachians from Siam and Indo-China (No. 2). Journal of the Natural History
Society of Siam, London, 6, 47–53.
Smith, M.A. (1940) The amphibians and reptiles obtained by Mr. Ronald Kaulback in Upper Burma. Records of the Indian
Museum, 42, 465–486.
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.
Swofford, D.L. (2001) PAUP*.Phylogenetic Analysis Using Parsimony (* and Other Methods), version 4. Sinauer Associates,
Sunderland, Massachusetts.
Taylor, E.H. (1962) New oriental reptiles. The University of Kansas Science Bulletin, 43, 209–263.
Theobald, W. (1876) Descriptive catalogue of the reptiles of British India. Thacker, Spink & Co., Calcutta, xiii, 238 pp.
Teynié, A. & David, P. (2010) Voyages naturalistes au Laos. Les Reptiles. Editions Revoir, Nohanent, France, 315 pp.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: Flexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876−4882.
Uetz, P. & Hosek, J. (Eds.) (2014) The Reptile Database. Available from: http://www.reptile-database.org (accessed 1 April
2014)
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.
http://dx.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.
http://dx.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.
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APPENDIX. Comparative material examined.
Cyrtodactylus cattienensis. Vietnam: Dong Nai Province: Cat Tien: IEBR A.0856 (holotype), IEBR 656, IEBR A.0855, ZFMK
88090–88095 (paratypes), and Vinh Cuu: IEBR A.0854, VNUH 2008.09.07 (paratypes); Ba Ria – Vung Tau Province: Nui
Dinh: IEBR A.0843–A.0845, VNUH 2008.0520 (paratypes).
Cyrtodactylus huongsonensis. Vietnam: Hanoi: Huong Son: IEBR A.2011.3 (holotype), ZFMK 92293 (paratype).
C. jaegeri. Laos: Khammuane Province, Thakkek: IEBR A.2013.55 (holotype) and NUOL R-2013.1 (paratype).
C. kingsadai. Vietnam: Phu Yen: Dai Lanh: IEBR A.2013.1 (holotype), IEBR A.2013.2–A.2013.3, ZFMK 94042–94043
(paratype).
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. pseudoquadrivirgatus. Vietnam: Thua Thien – Hue: A Luoi: ZFMK 83895 (holotype), IEBR 377, 379, 381–383 (paratypes);
Quang Tri Province: Huong Hoa: IEBR 2260–2267 (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); Khammuane Province: Ban Na Than: IEBR
KM.2012.14–15.
C. wayakonei. Laos: Luang Nam Tha: Vieng Phoukha: IEBR A.2010.01 (holotype), ZFMK 91016, MTD 47731, NUOL 2010.1
(paratypes).
