(Squamata: Gekkonidae) from Phu Yen Province ...

Zootaxa 3686 (4): 432–446 www.mapress.com / zootaxa / Copyright © 2013 Magnolia Press

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3686.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:E76D2B14-ED20-4D69-9624-63866C0C640F

A new Cyrtodactylus (Squamata: Gekkonidae) from Phu Yen Province, southern Vietnam THOMAS ZIEGLER1,2,8, TRUNG MY PHUNG3, MINH DUC LE4,5,6 & TRUONG QUANG NGUYEN2,7 1

Cologne Zoo, Riehler Straße 173, D-50735, Cologne, Germany. E-mail: [email protected] Zoological Institute, University of Cologne, Zülpicher Strasse 47b, D-50674 Cologne, Germany 3 Dong Khoi 9A, Tam Hiep, Bien Hoa, Dong Nai Province, Vietnam. E-mail: [email protected] 4 Faculty of Environmental Sciences, Hanoi University of Science, Vietnam National University, 334 Nguyen Trai Road, Hanoi, Vietnam. E-mail: [email protected] 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 79th Street, New York, New York 10024 7 Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet St., Hanoi, Vietnam. E-mail: [email protected] 8 Corresponding author. E-mail: [email protected] 2

Abstract We describe a new species of the genus Cyrtodactylus based on five adult specimens from Dai Lanh Cape, Tuy Hoa District, Phu Yen Province, southern Vietnam. Cyrtodactylus kingsadai sp. nov. is distinguished from the remaining Indochinese bent-toed geckos by a combination of the following characters: maximum SVL of 94 mm; dorsal pattern consisting of a dark nuchal loop, continuous or partly interrupted neck band and four in part irregular transverse body bands between limbs; internasal single; dorsal tubercles in 17–23 irregular transverse rows; ventrals in 39–46 longitudinal rows at midbody; lateral skin folds present, without interspersed tubercles; precloacal pores 7–9 plus in total 3–7 femoral pores in males (1-4 femoral pores on each side) with precloacal and femoral pore series separated from each other by 7–9 poreless scales; enlarged femoral scales and precloacal scales present; postcloacal spurs three; subcaudal scales transversely enlarged. This is the 29th species of Cyrtodactylus known from Vietnam. Key words: Cyrtodactylus kingsadai sp. nov., Phu Yen Province, southern Vietnam, morphology, phylogeny, taxonomy

Introduction Cyrtodactylus is the most diverse genus of gekkonids to date (e.g., Kluge 2001; Uetz 2013). Its widespread radiation occurs throughout tropical South Asia, Indochina, the Philippines, the Indo-Australian Archipelago, and the Solomon Islands in the East (Bauer & Henle 1994). Vietnam has been one of the countries with the most numerous discoveries of new Cyrtodactylus to date. Until 1997, only three species had been recorded for the country, C. condorensis (Smith), C. intermedius (Smith), and C. irregularis (Smith). Since then 25 additional species have been described from Vietnam, namely C. badenensis Nguyen, Orlov & Darevsky, C. bichnganae Ngo & Grismer, C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. bugiamapensis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, C. cucphuongensis Ngo & Chan, C. eisenmanae Ngo, C. grismeri Ngo, C. hontreensis Ngo, Grismer & Grismer, C. huongsonensis Luu, Nguyen, Do & Ziegler, C. huynhi Ngo & Bauer, C. martini Ngo, C. nigriocularis Nguyen, Orlov & Darevsky, C. paradoxus (Darevsky & Szczerbak), C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, C. phuquocensis Ngo, Grismer & Grismer, C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, C. takouensis Ngo & Bauer, C. thochuensis Ngo & Grismer, C. yangbayensis Ngo & Chan, and C. ziegleri Nazarov, Orlov, Nguyen & Ho (Ziegler et al. 2002;

432 Accepted by A. Bauer: 24 Jun. 2013; published: 15 Jul. 2013

Nguyen et al. 2006; Heidrich et al. 2007; Hoang et al. 2007; Orlov et al. 2007; Nazarov et al. 2008; Ngo 2008; Ngo & Bauer 2008; Ngo et al. 2008; Rösler et al. 2008; Geissler et al. 2009; Ngo & Chan 2010; Ngo & Grismer 2010; Ngo et al. 2010; Ziegler et al. 2010; Luu et al. 2011; Ngo 2011; Ngo & Chan 2011). Recent field research in southern Vietnam led to the discovery of more new populations of Cyrtodactylus. Based on morphological examination of five adult specimens, we herein describe a new species from Phu Yen Province, southern Vietnam.

Material and methods Sampling. Specimens of the new Cyrtodactylus were collected in coastal shrub vegetation intermixed with granite boulders in Dai Lanh Cape, Tuy Hoa District, Phu Yen Province by T. M. Phung between 2 and 25 September 2011 and from 19 to 20 January 2013. Specimens were anaesthetized, ethanol-fixed and subsequently deposited in the collections of the Institute of Ecology and Biological Resources (IEBR), Vietnam Academy of Science and Technology, Hanoi, Vietnam, and the Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Bonn, Germany. Molecular data and phylogenetic analyses. We used the protocols of Le et al. (2006) for DNA extraction, amplification, and sequencing. A fragment of the mitochondrial gene 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 Bayesian analysis (BA), as implemented in PAUP*4.0b10 (Swofford 2001) and MrBayes v3.2 (Huelsenbeck & Ronquist 2001), respectively. 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 HKY+I+G as selected by Modeltest v3.7 (Posada & Crandall 1998). Nodal support was evaluated using Bootstrap replication (BP) as calculated in PAUP and posterior probability (PP) in MrBayes v3.2. Uncorrected pairwise divergences were calculated in PAUP*4.0b10 (Table 1). TABLE 1. Samples used in molecular analyses; * = Cyrtodactylus pulchellus sensu lato (before the revision of Grismer et al. 2012). Species

Genbank No

Locality

Voucher information

Cyrtodactylus bugiamapensis Cyrtodactylus bugiamapensis Cyrtodactylus caovansungi Cyrtodactylus caovansungi Cyrtodactylus caovansungi Cyrtodactylus consobrinus Cyrtodactylus consobrinus Cyrtodactylus elok Cyrtodactylus elok Cyrtodactylus kingsadai sp. nov. Cyrtodactylus paradoxus Cyrtodactylus pubisulcus Cyrtodactylus pulchellus s.l.* Cyrtodactylus pulchellus s.l.* Cyrtodactylus quadrivirgatus Cyrtodactylus quadrivirgatus Cyrtodactylus ziegleri Cyrtodactylus ziegleri Cyrtodactylus ziegleri Cyrtodactylus cf. ziegleri

HM425553 HM888459 KF219678 KF219679 KF219680 HQ967204 HQ967205 HM888478 HM888479 KF188432 HM888464 HQ967199 HQ967202 HQ967203 HM888465 HM888466 HQ967210 HQ967211 HQ967212 HQ967213

Vietnam, Binh Phuoc Province Vietnam, Binh Phuoc Province Vietnam, Ninh Thuan Province Vietnam, Ninh Thuan Province Vietnam, Ninh Thuan Province Malaysia Malaysia Malaysia Malaysia Vietnam, Khanh Hoa Province Cambodia, Koh Tang Island Malaysia Malaysia Malaysia Malaysia Malaysia Vietnam, Dak Lak Province Vietnam, Dak Lak Province Vietnam, Dak Lak Province Vietnam, Dak Lak Province

NAP-00360 ZMMU R-13093-2 ITBCZ908 ITBCZ932 ITBCZ91113 ZMMU R-12644-1 ZMMU R-12644-2 ZMMU RAN 1991 ZMMU RAN 1992 IEBR A.2013.3 ZMMU RAN 1987 ZMMU R-13091-3 ZMMU R-12643-3 ZMMU R-12643-4 ZMMU RAN 1989 ZMMU RAN 1990 ZMMU R-13116-3 ZMMU R-13116-4 ZMMU R-13116-5 ZMMU R-13116-6

Morphological characters. For measurements and scale counts see Ziegler et al. (2010), terminology followed Rösler (1995). Measurements were taken with a slide-calliper (to the nearest 0.1 mm). Abbreviations used are: snout vent length (SVL), from tip of snout to vent; tail length (TaL), from vent to tip of tail; maximum head height (HH), from occiput to underside of jaws; head length (HL), from tip of snout to the posterior margin of the

A NEW CYRTODACTYLUS FROM SOUTHERN VIETNAM

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retroarticular; maximum head width across temporal region (HW); greatest diameter of orbit (OD); snout to eye distance (SE), measured from tip of snout to anterior corner of eye. The following scale counts were taken: supralabials (SL); infralabials (IL); nasal scales surrounding nare, from rostral to labial (but except counting rostral and labial), i.e. nasorostral, supranasal, postnasals (N); postrostrals or internasals (IN); postmentals (PM); dorsal tubercle rows (DTR), granular scales surrounding dorsal tubercles (GST); ventral scales in longitudinal rows at midbody between ventrolateral body folds (V); number of scales along the midbody from mental shield to anterior edge of cloaca (SLB); enlarged femoral scales (EFS); femoral pores (FP); precloacal pores (PP); postcloacal tubercles (PAT); subdigital lamellae on fourth finger (LD4); subdigital lamellae on fourth toe (LT4). Bilateral scale counts were given as left/right.

Results Phylogenetic analyses. The combined matrix contained 647 aligned characters. MP analysis of the dataset recovered two most parsimonious trees with 647 steps (CI = 0.60; RI = 0.75). One of the two trees is shown in Fig. 1. Sixty nine percent of the major nodes in the parsimonious tree (Fig. 1) received strong support (Bootstrap value ≥ 70%) (Hillis & Bull 1993) and the new species is placed with strong support in the clade containing C. bugiamapensis, C. caovansungi, C. consobrinus, C. paradoxus, C. pubisulcus, C. quadrivirgatus, and C. ziegleri (Bootstrap value = 72). In the Bayesian analysis, -lnL scores reached stationarity after 9,000 generations in both runs. The Bayesian topology is almost identical to the tree topology shown in Fig. 1 in major nodes (Fig. 1). The new species is significantly divergent from others in terms of genetic distance with the minimum pairwise divergence of approximately 13% in the mitochondrial fragment of COI (Table 2). Based on molecular comparisons and morphological distinctiveness with geographically close or morphologically similar species (e.g., C. caovansungi collected by T. M. Phung in January 2011 from the type locality in Nui Chua National Park, Ninh Thuan Province [IEBR A.2013.4] and C. yangbayensis [VNMN 03373, 03375] collected by C. V. Hoang in December 2011 from Hon Ba Nature Reserve, Dien Khanh District, Khanh Hoa Province, Vietnam) we came to the conclusion that the recent Cyrtodactylus collection from Dai Lanh represents an unnamed species which is described in the following as new:

FIGURE 1. One of the two most parsimonious maximum parsimony trees based on the partial COI gene. Number above and below branches are bootstrap values (>50%) and Bayesian posterior probabilities, respectively. Cyrtodactylus pulchellus = C. pulchellus sensu lato (before the revision of Grismer et al. 2012).

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435

Cyrtodactylus kingsadai sp. nov. Holotype. IEBR A.2013.1, adult male, collected on 25 September 2011 by T. M. Phung in coastal shrub vegetation intermixed with granite boulders in Dai Lanh Cape (12o55’N, 109o24’E), Tuy Hoa District, Phu Yen Province, southern Vietnam at an elevation between 50–100 m a.s.l. (Figs. 2–4). Paratypes. ZFMK 94042, adult male, IEBR A.2013.2 and ZFMK 94043, adult females, the same collection data as for the holotype (Fig. 5), and IEBR A.2013.3, adult male, collected on 20 January 2013 by T. M. Phung at the same locality. Diagnosis. A medium-sized Cyrtodactylus with a maximum SVL of 94 mm, distinguished from all congeners by the combination of the following characters: 1) dorsal pattern consisting of a dark nuchal loop, continuous or partly interrupted neck band and four in part irregular transverse body bands between limbs; 2) internasal single; 3) dorsal tubercles in 17–23 irregular transverse rows; 4) ventrals in 39–46 longitudinal rows at midbody; 5) lateral skin folds present, without interspersed tubercles; 6) precloacal pores 7–9 plus in total 3–7 femoral pores in males (1-4 femoral pores on each side), precloacal and femoral pore series separated from each other by 7–9 poreless scales; 7) enlarged femoral scales and precloacal scales present; 8) postcloacal spurs three; and 9) subcaudal scales transversely enlarged. Description of holotype. Size medium (SVL 83 mm, TaL 106 mm, tail tip regenerated), distance from posterior corner of eye to anterior margin of ear including ciliaria 6.5 mm, maximum horizontal ear diameter 1.1 mm; for further measurements see Table 3. TABLE 3. Selected measurements and morphological characters of the types series of Cyrtodactylus kingsadai sp. nov.; m = male, f = female, measurements in mm, * = regenerated or broken tail, m = mean, min. = minimum, max. = maximum, s = standard deviation. IEBR A.2013.1 (holotype)

ZFMK 94042 (paratype)

IEBR A.2013.3 (paratype)

IEBR A.2013.2 (paratype)

ZFMK 94043 (paratype)

min.-max. (m ± s)

Sex

m

m

m

f

f

SVL

83

88

83.4

92

94

max. 94

TaL

106*

45*

104.2*

117

117

max. 117

HH

9.3

10.4

11.3

10.0

9.9

max. 11.3

HL

24.2

24.7

25.2

25.6

26.7

max. 26.7

HW

15.1

16.3

18.0

16.2

17.2

max. 18.0

OD

5.7

6.4

6.8

6.3

6.0

max. 6.8

SE

10.9

11.4

10.6

11.0

11.7

max. 11.7

SL

13/14

14/13

11/12

13/13

11/12

11–14 (12.6±1.07)

IL

11/10

9/10

9/11

10/10

9/9

9–11 (9.8±0.79)

N

4/4

4/4

4/4

4/4

4/4

4

IN

1

1

1

1

1

1

PM

2

2

2

2

2

2

DTR

19

19

17

23

23

17–23 (20.2±2.68)

GST

10 or 11

10

9

9 or 10

9 or 10

9–11 (9.75±0.71)

V

46

46

39

43

43

39–46 (43.4±2.88)

SLB

165

174

177

177

173

165–177 (173.2±4.92)

EFS

11/12

11/12

10/9

12/12

12/12

9–12

FP

½

4/3

3/3

0

0

0–4

PP

8

7

9

8

4

4–9

PAT

3/3

3/3

3/3

3/3

3/3

3

LD4

19/19

20/19

19/20

21/20

21/20

19–21 (19.8±0.79)

LT4

21/21

21/23

21/22

25/24

23/21

21–25 (22.2±1.48)


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FIGURE 2. Adult male holotype of Cyrtodactylus kingsadai sp. nov. (IEBR A.2013.1) in preserved state. Photos T. Ziegler

Rostral wider than high (RW 3.6 mm, RH 2.1 mm, RW/RH 1.7) with an inverse Y-shaped median suture; supralabials 13/14; infralabials 11/10; nares bordered by rostral anteriorly, first supralabial laterally and four nasals posteriorly; supranasals separated from each other by two nasorostrals and a pentagonal internasal; medial snout scales slightly granular, those in contact with and nearby supralabials, flattened and larger than medial scales; upper anterior ciliaries three times larger than posterior ciliaries; head scales granular, smaller than median snout scales; back of head and temporal region with elongated to rounded, somewhat conical tubercles, 3–6 times larger than surrounding scales; mental triangular, slightly wider than rostral; one pair of enlarged postmentals, longer than wide, in broad contact posteriorly; postmentals bordering mental anteriorly, first two labials, one pair of distinctly enlarged gular scales, which are separated from each other by three small gular scales; dorsal scales somewhat granular to flattened, as large as medial snout scales; dorsal tubercles round, conical, surrounded by 10–11 granular scales, tubercles forming approximately 19 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2–3 times larger than dorsal granules, 46 longitudinal rows at midbody; lateral folds present, but not well developed, without interspersed tubercles; upper and lower arm with few slightly developed tubercles; dorsal hind



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limb covered by distinctly developed, flat to conical tubercles; series of distinctly enlarged femoral scales; femoral pore-bearing scales 11/12, distinctly enlarged, separated from precloacal, pore-bearing scales by 9 poreless femoral scales; femoral pores 1/2, precloacal pores 8, in an angular series; fingers and toes lacking distinct webbing; lamellae under first finger 16/16, under fourth finger 19/19, under first toe 15/16, and under fourth toe 21/21; claws surrounded by a small scale on upper and a large scale on lower sides; precloacal region covered by a patch of approximately 20 enlarged scales below precloacal, pore-bearing scales; precloacal groove absent; postcloacal tubercles 3/3, enlarged, on lateral surface of slight hemipenial swelling; dorsum of tail bearing distinct tubercles at base only, last 18 mm regenerated; subcaudals distinctly transversely enlarged, flat, smooth.

FIGURE 3. Head of the preserved holotype of Cyrtodactylus kingsadai sp. nov. (IEBR A.2013.1). Photos T. Ziegler

Coloration in ethanol. Ground colour light brownish-grey, with dark brownish-black dorsal pattern; dorsal surface of head with irregular dark markings, largest at occiput; a light canthal stripe extending from nostril to upper margin of eye, below and above framed by dark; a dark postocular streak, continuing to contact a somewhat irregularly shaped broad nuchal loop, with darker borders; postocular streak and nuchal loop bordered by thin light line; neck with a dark transverse band, also with dark borders, broadest in the centre; four more distinct dark transverse bands between limbs; dark transverse body bands somewhat irregularly shaped, with dark borders, and lighter vertebral region; interspaces between dark dorsal bands with irregular dark reticulation or blotches; dark tubercles in the nuchal loop and body bands, whereas light tubercles comprise the interspaces; lower flanks dark with small light blotches; upper surfaces of limbs with dark bands and reticulations; dorsal surface of tail with approximately ten broad, dark transverse bands; light interspaces brownish-grey at tail base, more greyish-white towards the regenerated tail tip; sides of dark transverse tail bands with small light blotches; gular region yellowish-cream with indistinct grey marbling; venter cream with light grey belly; tail surface light to dark grey towards the tip. Variation of paratypes. For the variation in scalation see Table 3, and for colour pattern variation see Fig. 5. In the male paratypes, the precloacal pore-bearing scales are separated by 7-9 poreless scales from the femoral pore bearing scales. The neck band is somewhat medially interrupted in the male paratype IEBR A.2013.3, and the dark transverse body bands are slightly interrupted medially in the female paratype ZFMK 94043. With respect to


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sexual dimorphism, the female paratypes are larger, lack hemipenial swellings at the tail base, and the femoral pores are absent.

FIGURE 4. Cloacal region of the preserved holotype of Cyrtodactylus kingsadai sp. nov. (IEBR A.2013.1). Photo T. Ziegler

Comparisons. Comparisons are based on the original descriptions or descriptions provided in broader faunal and taxonomic publications (e.g., Grismer et al. 2008; Rösler & Glaw 2008; Bauer et al. 2009; 2010; Ngo & Grismer 2010; Ngo & Pauwels 2010; Sumontha et al. 2010; Ziegler et al. 2010; David et al. 2011; Iskandar et al. 2011; Luu et al. 2011; Ngo 2011; Ngo & Chan 2011; Schneider et al. 2011; Nazarov et al. 2012; Ngo & Grismer 2012). The most important morphological distinguishing characters between Cyrtodactylus kingsadai sp. nov. and its Vietnamese congeners are summarized in Table 4. Furthermore, Cyrtodactylus kingsadai sp. nov. has three postcloacal tubercles in males and thus differs from C. bichnganae, which has only two postcloacal tubercles in males; C. bichnganae also has 28 femoral and precloacal pores separated by a short diastema on each side. From the similar C. caovansungi, Cyrtodactylus kingsadai sp. nov. differs by having 9–12 (vs. 8) enlarged femoral scales on each side, by 17–23 (vs. 16–18) dorsal midbody tubercle rows, and by 5 (4 + 1) versus 4 dark transverse dorsal bands (except for nuchal loop). Furthermore, Cyrtodactylus kingsadai sp. nov. lacks tubercles on the lateral skin fold and on the dorsal tail surface and thus differs from C. condorensis; in addition, the latter species only has precloacal pores (4–7) and a blotched dorsal pattern. Cyrtodactylus kingsadai sp. nov. has precloacal scales in males which are lacking in C. cucphuongensis. Cyrtodactylus kingsadai sp. nov. has 3–7 femoral pores in total in males and thus differs from C. huongsonensis (17). Cyrtodactylus kingsadai sp. nov. has precloacal pores separated from femoral pores in males, which are a contiguous series in C. phongnhakebangensis and C. roesleri. Cyrtodactylus kingsadai sp. nov. differs from C. yangbayensis by having precloacal pores also in females, by the presence of a distinct broad dark nuchal loop which is broken or V-shaped in C. yangbayensis, and by distinct broad versus more numerous irregular rows of narrow dark transverse dorsal body bands; furthermore, males of C. yangbayensis usually lack femoral pores (of four males, only one individual had 2 femoral pores) versus 1–4 femoral pores present on each side in Cyrtodactylus kingsadai sp. nov.



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With respect to the remaining Cyrtodactylus (except for zoogeographically distant species from the PapuaAustralian region, the Solomon Islands, India, Sri Lanka, the Nicobar islands and Nepal), Cyrtodactylus kingsadai sp. nov. has transversely enlarged subcaudals and thus differs from the following species which lack enlarged subcaudals: C. aequalis Bauer, C. agusanensis (Taylor), C. annulatus (Taylor), C. batucolus Grismer, Chan, Grismer, Wood & Belabut, C. brevidactylus Bauer, C. buchardi David, Teynié & Ohler, C. cavernicolus Inger & King, C. fumosus (Müller), C. gansi Bauer, C. halmahericus Mertens, C. jambangan Welton, Siler, Diesmos & Brown, C. jellesmae (Boulenger), C. lateralis (Werner), C. majulah Grismer, Wood & Lim, C. malayanus (De Rooij), C. mandalayensis Mahony, C. marmoratus Gray, C. matsuii Hikida, C. nuaulu Oliver, Edgar, Mumpuni, Iskandar & Lilley, C. pantiensis Grismer, Chan, Grismer, Wood & Belabut, C. payacola Johnson, Quah, Anuar, Muin, Wood, Grismer, Greer, Onn, Ahmad, Bauer & Grismer, C. philippinicus (Steindachner), C. pubisulcus Inger, C. quadrivirgatus Taylor, C. semenanjungensis Grismer & Leong, C. seribuatensis Youmans & Grismer, C. stresemanni Rösler & Glaw, C. sworderi (Smith), C. tautbatorum Welton, Siler, Diesmos & Brown, C. tiomanensis Das & Lim, C. wakeorum Bauer, C. wetariensis (Dunn), C. yoshii Hikida, and C. zhaoermii Shi & Zhao.

FIGURE 5. a–c: Adult male paratype (ZFMK 94042), d: uncollected adult female 2.9.2011, and e–f: adult female paratype (IEBR A.2013.2) of Cyrtodactylus kingsadai sp. nov. in life. Photos T. M. Phung.



441

Cyrtodactylus kingsadai sp. nov. has femoral pores in males and thus differs from the following species which lack femoral pores: C. angularis (Smith), C. aurensis Grismer, C. ayeyarwadyensis Bauer, C. batik Iskandar, Rachmansah & Umilaela, C. chanhomeae Bauer, Sumontha & Pauwels, C. chrysopylos Bauer, C. consobrinoides (Annandale), C. deveti (Brongersma), C. elok Dring, C. feae (Boulenger), C. ingeri Hikida, C. jarujini Ulber, C. khasiensis (Jerdon), C. oldhami (Theobald), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, C. papilionoides Ulber & Grossmann, C. peguensis (Boulenger), C. rubidus (Blyth), C. sanook Pauwels, Sumontha, Latinne & Grismer, C. sumonthai Bauer, Pauwels & Chanhome, and C. variegatus (Blyth). Cyrtodactylus kingsadai sp. nov. has 7-9 precloacal pores plus in total 3–7 femoral pores in males, which are separated by poreless scales and thus differs from the following species which have a contiguous series of precloacal-femoral pores: C. astrum Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. australotitiwangsaensis Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. bintangrendah Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. bintangtinggi Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. langkawiensis Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. lekaguli Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels, C. lomyenensis Ngo & Pauwels (32–40), C. phuketensis Sumontha, Pauwels, Kunya, Nitikul, Samphantamit & Grismer (33-36), C. surin Chan-Ard & Makchai (34), C. tamaiensis Mahony (40), and C. trilatofasciatus Grismer, Wood, Quah, Anuar, Muin, Sumontha, Ahmad, Bauer, Wangkulangkul, Grismer & Pauwels. Cyrtodactylus kingsadai sp. nov. has 7–9 precloacal pores in males and 4–8 precloacal pores in females and thus differs from the following species which have distinctly higher precloacal pore counts: C. annandalei Bauer (11–12), C. baluensis (Mocquard) (9–11), C. durio Grismer, Anuar, Quah, Muin, Onn, Grismer & Ahmad (12), C. interdigitalis Ulber (14), C. russelli Bauer (15), C. spinosus Linkem, McGuire, Hayden, Setiadi, Bickford & Brown (12–13), and C. teyniei David, Nguyen, Schneider & Ziegler (14 in the single known specimen, an adult female). The following Cyrtodactylus species differ from Cyrtodactylus kingsadai sp. nov. by the absence of precloacal and femoral pores in both sexes: C. darmandvillei (Weber, 1890), C. gordongekkoi (Das) (see Biswas 2007), C. jarakensis Grismer, Chan, Grismer, Wood & Belabut, C. laevigatus (Darevsky), C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome, and C. wallacei Hayden, Brown, Gillespie, Setiadi, Linkem, Iskandar, Umilaela, Bickford, Riyanto, Mumpuni & McGuire. Cyrtodactylus kingsadai sp. nov. has 39–46 ventral scales at midbody and thus differs from C. agamensis (Bleeker) (67), C. consobrinus (Peters) (65–70), C. erythrops Bauer, Kunya, Sumontha, Niyomwan, Panitvong, Pauwels, Chanhome & Kunya (28), C. gubaot Welton, Siler, Linkem, Diesmos & Brown (54–67), C. leegrismeri Chan & Norhayati (27–35), C. macrotuberculatus Grismer & Norhayati (19–22), C. mamanwa Welton, Siler, Linkem, Diesmos & Brown (57–70), C. pulchellus Gray (33–35), C. slowinskii Bauer (27–32), C. sumuroi Welton, Siler, Linkem, Diesmos & Brown (53–58), C. tigroides Bauer, Sumontha & Pauwels (34), and C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler (31–35). Cyrtodactylus kingsadai sp. nov. has lateral skin folds without interspersed tubercles and thus differs from C. brevipalmatus (Smith), and C. mimikanus (Boulenger). Cyrtodactylus kingsadai sp. nov. has 17–23 dorsal tubercle rows and thus differs from C. redimiculus King (14–16), which also lacks a ventro-lateral fold. Cyrtodactylus kingsadai sp. nov. differs from C. auribalteatus Sumontha, Panitvong & Deein by different dorsal pattern (dorsal pattern consisting of only three dark bands between limb insertions), from C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya in having internasal. Distribution. Cyrtodactylus kingsadai sp. nov. is currently known only from the type locality in Vietnam (Fig. 6). Etymology. The new species is named in memory of our cooperation partner and friend Phouthone Kingsada from the National University of Laos, Vientiane, with whom we have conducted several field excursions in the forests of Laos and who unfortunately died too young in December 2012. Ecological notes. The type series was found at night in coastal shrub vegetation intermixed with granite boulders at elevations between 50–100 m a.s.l. (Fig. 7). The predominant vegetation consists of small prickly shrubs representing species of the families Ebenaceae, Dipterocarpaceae, Annonaceae, and Fabaceae. Further specimens were sighted but not collected. Cyrtodactylus kingsadai co-occurs with Gekko truongi (voucher


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specimen from Dai Lanh: ZFMK 94044), which recently was described by Phung & Ziegler (2011) from Cuc Dong Cape, Ninh Hoa District, Khanh Hoa Province.

FIGURE 6. Map showing the type locality of Cyrtodactylus kingsadai sp. nov. in Phu Yen Province, Vietnam.



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FIGURE 7. Habitat of Cyrtodactylus kingsadai sp. nov. Photo T. M. Phung.

Discussion Our phylogenetic results show that the new species is not closely related to any of the compared Cyrtodactylus species. However, this could be an artifact of our data set, which does not have enough informative characters to resolve closer relationships. Phenetically, the new species resembles C. caovansungi from southeastern Vietnam (type locality Ninh Thuan Province), but besides genetic differences, it inhabits different ecosystem, such as the geographically close C. yangbayensis (type locality Khanh Hoa Province), which is a forest dweller as well (Orlov et al. 2007, Ngo & Chan 2010), whereas C. kingsadai sp. nov. inhabits open coastal shrub vegetation.

Acknowledgements T.M. Phung is grateful to T.L.T. Nguyen (University of Ulsan, South-Korea), T.K. Tran, N.T. Hoang, T.H.M. Vu, and H.V. Bui (Ho Chi Minh City) for assistance in the field. S.N. Nguyen and J. Che (Kunming Institute of Zoology) kindly sent us the sequences of C. caovansungi. For the loan of specimens, we acknowledge C.X. Le, C.T. Pham, C.V. Hoang (IEBR, Hanoi) and T.T. Nguyen (Vietnam National Museum of Nature, Hanoi), as well as for the kind permission to donate a type specimen to the herpetological collection of the ZFMK, Bonn, Germany. We also would like to thank A. Rauhaus (Cologne Zoo) for the kind arrangement of the colour plates, Nicole Schneider, Mona van Schingen and Martin Kutzner (Bonn and Cologne Universities) for help with morphological


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data collection, Duong Thuy Ha (Hanoi) for laboratory assistance, and E. Sterling (New York) and K. Koy (Berkeley) for providing the map. O.S.G. Pauwels (Brussels) and L.L. Grismer (Riverside) kindly revised a previous version of the manuscript. T.Q. Nguyen thanks M. Bonkowski (Cologne) for support of his work in Germany. Research of T.Q. Nguyen in Germany is funded by the Alexander von Humboldt Stiftung/Foundation (VIE 114344). M.D. Le was supported by the National Foundation for Science and Technology Development of Vietnam (NAFOSTED: Grant No. 106.15-2010.30). Equipment for field work in Vietnam was supported by the Cologne Zoo.

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