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A new species of Cyrtodactylus (Squamata: Gekkonidae)
from Ninh Binh Province, Vietnam
DZUNG TRUNG LE1, TRUONG QUANG NGUYEN2,8, MINH DUC LE3,4,5 & THOMAS ZIEGLER6,7
1

Faculty of Biology, Hanoi National University of Education, 136 Xuan Thuy Road, Hanoi, Vietnam.
E-mail:
2
Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi, Vietnam.
E-mail:
3
Faculty of Environmental Sciences, Hanoi University of Science, Vietnam National University, 334 Nguyen Trai Road, Hanoi,
Vietnam. E-mail:
4
Centre for Natural Resources and Environmental Studies, Hanoi National University, 19 Le Thanh Tong, Hanoi, Vietnam
5
Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024
6
AG Zoologischer Garten Köln, Riehler Strasse 173, D–50735 Cologne, Germany. E–mail:



Vietnam, in July and August 2015. Specimens were anaesthetized and fixed in approximately 85% ethanol, then
later transferred to 70% ethanol for permanent storage. Specimens referred to in this paper are deposited in the
collections of the Museum of Biology, Hanoi National University of Education (HNUE), Hanoi; the Institute of
Ecology and Biological Resources (IEBR), Hanoi; the Tay Bac University (TBU), Son La Province; Muséum
d'histoire naturelle, Geneva (MHNG), Switzerland; the Vietnam Forest Museum (VFM), Forest Inventory and
Planning Institute, Hanoi; the Vietnam National Museum of Nature (VNMN), Hanoi; the Vietnam National
University of Forestry (VUF), Hanoi, Vietnam; the Namlik Ecovillage Museum (NEM), Ban That Wang Monh,
Vientiane Province, Laos; the National University of Laos (NUOL), Vientiane, Laos; the Senckenberg
Naturhistorische Sammlungen Dresden, Museum für Tierkunde (MTD), Dresden; and the Zoologisches
Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany.
Molecular data and phylogenetic analyses. We sequenced two samples of Cyrtodactylus collected from Van
Long Wetland Nature Reserve and the holotype of C. huongsonensis (IEBR A.2011.3A, see Luu et al. 2011). We
also included all available sequences from members of the C. wayakonei species group (Fig. 1, Table 1). The
species group was defined as clade B in Nguyen et al. (2015). Two species, C. elok Dring and C. pulchellus Gray
(see Grismer et al. 2012), 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´107. 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 11 in the
Bayesian analysis, as -lnL scores reached stationarity after 11,000 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 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

The final matrix consisted of 657 aligned characters, of which 212 are parsimony informative. The alignment
contained no gap. MP analysis of the dataset recovered two most parsimonious trees with 578 steps (CI = 0.58; RI
= 0.76). In the ML analysis, the score of the single best tree found was 3361.12 after 2448 arrangements were tried.
The topology derived from the Bayesian analysis (Fig. 1) is similar to clade B in Nguyen et al. (2015), but C.
bichnganae was supported as the sister taxon to C. huongsonensis + the new species instead of being placed as the
basal taxon of the clade as shown in Nguyen et al. (2015) (Fig. 1). However, both placements received low
statistical support values from all analyses. The new species was recovered as the sister taxon to C. huongsonensis
with high statistical support values from all phylogenetic analyses. These two species were approximately 5.0–
5.5% genetically divergent from each other (Table 1).
TABLE 1. Uncorrected (“p) distance matrix showing percentage pairwise genetic divergence (COI) between new and
closely related species
Species name

1

2

3

1. C. bichnganae
(KT004372)

–

2. C. bobrovi
(KT004367-8)

16.3

–

3.8

17.8

15.4

–

6. C. puhuensis
(KF929529)

18.9

7.3–7.5

17.8

18.3

7.4

7. Cyrtodactylus
soni sp. nov.
(KX430032-3)

13.4–13.9 16.7–16.9 16.4–16.7 5.0–5.5

8. C. spelaeus
(KP199947-8)



16.1–16.4 19.1–19.5 –

9.4

10.4

15.7–16.1

16.6–16.9 18.1–18.4 18.2–18.4 17.2–17.7

–
11.7–12.0 –
16.1–16.5 15.8–16.2 –

Cyrtodactylus soni sp. nov.
(Figs. 2–4)
Holotype. HNUE VL.2015.78, adult male, collected on 24 July 2015 by D. T. Le, A. M. Luong, D. T. Pham, and
N. H. Nguyen, in the karst forest near Da Han Village (20o25.067’N, 105o51.467’E, elevation 17 m a.s.l.), Gia Hoa
Commune, within Van Long Wetland Nature Reserve, Gia Vien District, Ninh Binh Province, Vietnam.
Paratypes. IEBR R.2016.4, adult female, HNUE VL.2015.94, adult female, collected on 24 July 2015; IEBR
R.2016.5, adult male, HNUE VL.2015.131, adult female, and HNUE VL.2015.132, adult female, collected on 18
August 2015, the same data as the holotype.
Diagnosis. The new species can be distinguished from other members of the genus Cyrtodactylus from
Indochina by a combination of the following characters: medium size (SVL up to 103 mm); internasal single;
dorsal tubercles in 10–13 irregular rows; ventral scale rows 41–45; lateral skin folds present, without interspersed

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broadened lamellae), finger II 14 (4), finger III 15 (4), finger IV 16 (4), finger V 16 (5), toe I 11 (2), toe II 14 (4),
toe III 17 (5), toe IV 18 (6), toe V 19 (7).
Tail regenerated (TaL 70.6 mm); postcloacal spurs 2/2; dorsal tail base with tubercles; subcaudals distinctly
enlarged.
Coloration in life. Ground color light brownish grey; dorsal surface of head with irregular dark markings,
largest at occiput; a dark postocular streak, edged in yellowish white, in contact with nuchal loop; neck with a dark
transverse band; dorsum with five distinct dark transverse bands between limb insertions, edged by yellowish
tubercles; upper surfaces of limbs with dark bands and reticulations; dorsal surface of the regenerated tail with
indistinct transverse bands; gular region cream with indistinct grey marbling; venter cream; ventral surface of the
regenerated tail dark grey with yellow marbling.
Morphological comparisons. We compared the new species with its congeners from Vietnam and
neighbouring countries in mainland Indochina, including Laos, Cambodia, and Thailand based on the examination
of specimens (see Appendix) and data obtained from the literature (Bauer et al. 2002, 2003, 2010; David et al.
2004, 2011; Geissler et al. 2009; Hoang et al. 2007; Kunya et al. 2014, 2015; Luu et al. 2011, 2014, 2015, 2016a,
2016b, 2016c; Nazarov et al. 2008, 2012, 2014; Ngo 2011, 2013; Ngo & Grismer 2010; Ngo & Chan 2011;
Nguyen et al. 2010, 2015; Nguyen et al. 2013, 2014; Panitvong et al. 2014 ; Pauwels & Sumontha 2014; Pauwels
et al. 2013, 2014a, 2014b, 2016; Phung et al. 2014; Schneider et al. 2011, 2014 a, 2014b; Smith 1921; Sumontha et
al. 2015; Ziegler et al. 2010, 2013).
Cyrtodactylus soni sp. nov. has distinctly enlarged subcaudals, which are only slightly or not enlarged in the
following species: C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. buchardi
David, Teyni & Ohler, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C.
cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, C. cryptus Heidrich, Rösler, Vu, Böhme
& Ziegler, C. cucdongensis Schneider, Phung, Le, Nguyen & Ziegler, C. huynhi Ngo & Bauer, C. irregularis
(Smith), C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. phuocbinhensis Nguyen, Le, Tran, Orlov,


LE ET AL.


FIGURE 2. The male holotype (HNUE VL.2015.78) of Cyrtodactylus soni sp. nov. in life (A: Dorsal view, B: Ventral view).
Photo D. T. Le.

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FIGURE 3. Cloacal (A) and femoral (B) regions of the holotype (HNUE VL.2015.78) of Cyrtodactylus soni sp. nov. in
preservative. Photo D. T. Le.

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LE ET AL.


FIGURE 4. Female paratype (IEBR R.2016.4) of Cyrtodactylus soni sp. nov. in life: A) Dorsal view and B) Ventral view.
Photo D. T. Le.

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275

F

F

F

F

SVL

89.5

89.2

103

88.7

89.5

98.5

88.7–103

TaL

70.6*

82.8*


18.7

19

17.3

17.7

19.3

17.3–19.3

HH

11.2

11

11

10

10

11.1

10–11.2

OD


EE

8.3

7

8.6

7.9

7.5

8.4

7–8.6

NE

8.7

8.4

8.7

8.8

8.3

9.1


16.3

13.8–16.3

CrusL

17.5

17.2

18.8

17.2

16.7

19.5

16.7–19.5

AG

39.8

36

47.4

37.9


2.9

3.2

3.3

2.9–3.8

IOD

6.8

7.4

6

7.4

6.7

7.5

6–7.5

SPL

10/10

11/10


5/4

4/4

4/4

4/4

3/3

3–5

IN

1

1

1

1

1

1

1

CIL


GST

9 or 10

9–11

9 or 10

9–11

9–10

9–11

9–11

V

41

42

45

45

44

43


8/6

5–8

PP

6

7

7 (pitted scales)

8 (pitted scales)

7 (pitted scales)

7 (pitted scales)

6–8

PAT

2/2

3/3

3/2

2/2


9/9

11/11

9/10

8–11

I

2+9

3+10

3+10

3+8

2+10

2+10

11–13

II

4+10

5+12


5+14

5+10

5+10

5+14

6+12

15–19

V

5+11

4+11

6+12

4+11

4+12

4+11

15–18

2+9


14–18

III

5+12

6+14

7+14

5+14

5+15

7+12

19–21

IV

6+12

7+13

9+13

9+12

7+15


Grismer & Pauwels (31–38), C. bansocensis Luu, Nguyen, Le, Bonkowski & Ziegler (34), C. calamei Luu,
Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (35–39), C. chanhomeae Bauer, Sumontha & Pauwels (32), C.
darevskii Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov (38–44), C. erythrops
Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya (10+9+9), C. hinnamnoensis Luu,
Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (36–44), C. jaegeri Luu, Calame, Bonkowski, Nguyen &
Ziegler (44), C. jarujini (52–54), C. khammouanensis Nazarov, Poyarkov, Orlov, Nguyen, Milto, Martynov,
Konstantinov & Chulisov (40–44), C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer,
Wangkulangkul, Grismer & Pauwels (31–41), C. lomyenensis Ngo & Pauwels (39–40), C. multiporus (58–60), C.
phongnhakebangensis Ziegler, Rösler, Herrmann & Vu (32–42), C. rufford Luu, Calame, Nguyen, Le, Bonkowski
& Ziegler (42–43), C. sommerladi Luu, Bonkowski, Nguyen, Le, Schneider, Ngo & Ziegler (20–26), and C.
soudthichaki (29).
Cyrtodactylus soni sp. nov. has a dorsum with banded color pattern, which is blotched in C. brevipalmatus
(Smith), C. buchardi, C. bugiamapensis, C. erythrops, C. irregularis, C. jarujini, C. phetchaburiensis, C.
phuocbinhensis, C. pseudoquadrivirgatus, C. taynguyenensis, C. teyniei David, Nguyen, Schneider & Ziegler, and
C. thuongae; the dorsum is uniformly brown in C. nigriocularis and striped in C. oldhami, C. quadrivirgatus, and
C. ranongensis.
Cyrtodactylus soni sp. nov. differs from C. kingsadai Ziegler, Phung, Le & Nguyen by having fewer dorsal
tubercle rows (10–13 vs. 17–23 in C. kingsadai).
Cyrtodactylus soni sp. nov. differs from the following species by having more ventral scale rows (41–45): C.
badenensis (25–29), C. bichnganae (30–31), C. buchardi (30), C. chanhomeae (36–38), C. chauquangensis (36–
38), C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri (29–35), C.
erythrops (28), C. grismeri (33–38), C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu, Sirisamphan &
Pauwels (29–34), C. jaegeri (31–32), C. jarujini (32–38), C. khammouanensis (32–38), C. khelangensis Pauwels,

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Discussion
The phylogenetic analyses showed that C. soni together with C. huongsonensis form an independent and wellsupported clade within the C. wayakonei species group (sensu Nguyen et al. 2015; Luu et al. 2016a, 2016b, 2016c).
Although C. bichnganae was also clustered with the clade, its relationship with the other two species was only
weakly corroborated by all analyses. Biogeographically, C. soni and C. huongsonensis are distributed in the same
mountain range southeast of Hanoi. The localities of the two species are about 30 km apart from each other. A
similar level of species richness has been found in other groups of Cyrtodactylus in Vietnam and Laos (Nguyen et
al. 2015, Luu et al. 2014, 2016b, 2016c). It is therefore important to continue taxonomic research in the genus
because it likely harbors a high level of undescribed diversity, especially in karst habitats (Luu et al. 2016a).

Acknowledgements
We are grateful to the directorates of the Forest Protection Department of Ninh Binh Province, and Van Long
Wetland Nature Reserve for support of our field work and issuing relevant permits. D. T. Le thanks A.M. Luong,
D.T. Pham (Hanoi) and staff of Van Long Wetland Nature Reserve for their assistance in the field. We thank D.T.
Pham and H. T. Ngo (Hanoi) for laboratory assistance, E. Sterling (New York) and K. Koy (Berkeley) for providing
the map. Many thanks to S.N. Nguyen (Ho Chi Minh City), O.S.G. Pauwels (Brussels) and A. M. Bauer (Villanova)
for their helpful comments. For the fruitful cooperation within joint biodiversity research and conservation projects
we cordially thank T. H. Tran (IEBR, Hanoi), T. Pagel and C. Landsberg (Cologne Zoo). This research is funded by
the Vietnam National Foundation for Science and Technology Development (NAFOSTED) under grant number
106-NN.05-2014.34 and Cologne Zoo. Equipment was funded by the Alexander von Humboldt Foundation (VIE
1143441).

References
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.

A NEW CYRTODACTYLUS FROM VIETNAM

Zootaxa 4162 (2) © 2016 Magnolia Press ·


/>Luu, V.Q., Bonkowski, M., Nguyen, T.Q., Le, M.D., Schneider, N., Ngo, H.T. & Ziegler, T. (2016a) Evolution in karst massifs:
Cryptic diversity among bent-toed geckos along the Truong Son Range with descriptions of three new species and one new
country record from Laos. Zootaxa, 4107 (2), 101−140.
/>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.
/>Luu, V.Q., Calame, T., Nguyen, T.Q., Bonkowski, M. & Ziegler, T. (2015) A new species of Cyrtodactylus (Squamata:
Gekkonidae) from the limestone forest of Khammouane Province, central Laos. Zootaxa, 4058 (2), 388−402.
/>Luu, V.Q., Calame, T., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016b) Cyrtodactylus rufford, a new cavedwelling Bent-toed Gecko (Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4067 (2),
185−199.
/>Luu, V.Q., Nguyen, T.Q., Do, H.Q. & Ziegler, T. (2011) A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son
limestone forest, Hanoi, northern Vietnam. Zootaxa, 3129, 39–50.
Luu, V.Q., Nguyen, T.Q., Le, M.D., Bonkowski, M. & Ziegler, T. (2016c) A new species of karst-dwelling Bent-toed Gecko
(Squamata: Gekkonidae) from Khammouane Province, central Laos. Zootaxa, 4079 (1), 87−102.
/>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.A., Poyarkov, N.A., Orlov, N.L., Nguyen, S.N., Milto, K.D., Martynov, A.A., Konstantinov, E.L. & Chulisov, A.S.
(2014) A review of genus Cyrtodactylus (Reptilia: Sauria: Gekkonidae) in fauna of Laos with description of four new
species. Proceedings of the Zoological Institute, RAS, 318, 391–423.
Nazarov, R., Poyarkov, N.A., Orlov, N.I., 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. (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.

280 · Zootaxa 4162 (2) © 2016 Magnolia Press

LE ET AL.



Ba Den Mountain, Tay Ninh Province, southern Vietnam. Zootaxa, 3764 (3), 347−363.
/>Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817−818.
/>Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. &
Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model
space. Systematic Biology, 61, 539−542.
/>Schneider, N., Nguyen, T.Q., Le, M.D., Nophaseud, L., Bonkowski, M. & Ziegler, T. (2014a) A new species of Cyrtodactylus
(Squamata: Gekkonidae) from the karst forest of northern Laos. Zootaxa, 3835 (1), 80–96.
/>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. (2014b) A new Cyrtodactylus (Squamata: Gekkonidae) from
Khanh Hoa Province, southern Vietnam. Zootaxa, 3785 (4), 518–532.
/>Smith, M.A. (1920) Reptiles and batrachians collected on Pulo Condore. Journal of Natural History Society of Siam, 4, 93–97.
Sumontha, M., Pauwels, O.S.G., Panitvong, N., Kunya, K. & Grismer, L.L. (2015) A new lowland forest Bent-toed Gecko
(Squamata: Gekkonidae: Cyrtodactylus) from Ranong Province, peninsular Thailand. Zootaxa, 3911 (1), 106–118.
/>Swofford, D.L. (2001) PAUP*. Phylogenetic Analysis Using Parsimony (* and Other Methods), version 4. Sinauer Associates,

A NEW CYRTODACTYLUS FROM VIETNAM

Zootaxa 4162 (2) © 2016 Magnolia Press ·

281


Sunderland, Massachusetts.
Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: Flexible
strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876−4882.
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.