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A new (: ) from Huong Son limestone forest, Hanoi, northern Vietnam

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Zootaxa 3129: 39–50 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2011 · Magnolia Press ISSN 1175-5334 (online edition)

A new Cyrtodactylus (Squamata: Gekkonidae) from Huong Son limestone forest, Hanoi, northern Vietnam

VINH QUANG LUU1, TRUONG QUANG NGUYEN2, HUY QUANG DO1, THOMAS ZIEGLER3,4 1Department of Wildlife, Faculty of Natural Resource and Environmental Management, The Forestry University of Vietnam, Xuan Mai, Chuong My, Ha Noi, Vietnam 2Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet St., Hanoi, Vietnam 3Cologne Zoo, Riehler Straße 173, D-50735, Cologne, Germany 4Corresponding author. E-mail: [email protected]

Abstract

We describe a new of the genus Cyrtodactylus based on two adult specimens from Huong Son limestone forest, Hanoi, Vietnam. Cyrtodactylus huongsonensis sp. nov. is distinguished from the remaining Indochinese bent-toed geckos by a combination of the following characters: medium-sized, with a maximum SVL of 89.8 mm; dorsal pattern consisting of dark nuchal loop, neck band and five in part irregular transverse body bands between limbs; two enlarged lateral chin- shields in contact with first postmental pair; dorsal tubercles present on occiput, body, forearms, hind limbs and tail base; 14–16 irregularly running dorsal tubercle rows; ventrals in 41–48 longitudinal rows at midbody; lateral skin folds present, without interspersed tubercles; six precloacal pores plus in total 17 femoral pores in males, which are separated by 8–12 poreless scales; enlarged femoral scales and precloacal scales present; three postcloacal spurs in males; subcaudal scales transversely enlarged. This is the 24th species of Cyrtodactylus known from Vietnam.

Key words: Cyrtodactylus huongsonensis sp. nov., Hanoi, Vietnam, morphology,

Introduction

The genus Cyrtodactylus is the most diverse group of gekkonids to date (e.g., Kluge 2001, Uetz et al. 2011). The widespread radiation occurs throughout tropical South Asia, Indochina, the Philippines, the Indo-Australian Archi- pelago, and the Solomon Islands in the East (Bauer & Henle 1994). Vietnam has been one of the countries of the most numerous discoveries of new Cyrtodactylus to date. Until 1997, only three species had been recorded for the country, C. condorensis (Smith, 1921), C. intermedius (Smith, 1917), and C. irregularis (Smith, 1921). Since then 20 additional species have been described from Vietnam, namely C. badenensis Nguyen, Orlov & Darevsky, 2006, C. bichnganae Ngo & Grismer, 2010, C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, 2009, C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007, C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007, C. eisenmanae Ngo, 2008, C. grismeri Ngo, 2008, C. hontreensis Ngo, Grismer & Grismer, 2008, C. huynhi Ngo & Bauer, 2008, C. martini Ngo, 2011, C. nigriocularis Nguyen, Orlov & Darevsky, 2006, C. paradoxus (Darevsky & Szczerbak, 1997), C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, 2003, C. phuquocensis Ngo, Grismer & Grismer, 2010, C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, 2008, C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010, C. takouensis Ngo & Bauer, 2008, C. yangbayensis Ngo & Chan, 2010, and C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008 (Ziegler et al. 2002, 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, Ngo 2011). Recent field research in northern Vietnam led to the discovery of more new populations of Cyrtodactylus. Based on morphological examination of two adult specimens, we herein describe a new species from Huong Son limestone forest, My Duc District, Hanoi.

Accepted by A. Bauer: 2 Nov. 2011; published: 14 Dec. 2011 39 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Material and methods

Specimens of the new Cyrtodactylus were collected in Huong Son limestone forest by Vinh Quang Luu, Hieu Van Pham, and Nghia Van Ha on 13 June 2011. Specimens were anaesthetized, ethanol-fixed and subsequently depos- ited in the collections of the Institute of Ecology and Biological Resources, Vietnamese Academy of Science and Technology Hanoi (IEBR), Vietnam, and the Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK), Germany. For measurements and scale counts see Ziegler et al. (2010), terminology followed Rösler (1995). Measure- ments were taken with a slide-calliper (to the nearest 0.1 mm). Abbreviations are as follows: 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 retroarticular; max- imum head width (HW); greatest diameter of orbit (OD); snout to eye distance (SE), measured from tip of snout to anteriormost point of eye. Scale counts were taken as follows: 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); number of distinguishable eyelid fringe scales or ciliaria (CIL); postmentals (PM); dorsal tubercle rows (DTR), granular scales surrounding dorsal tubercles (GST); ventral scales in longitudinal rows at midbody (V); number of scales along the midbody from mental 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.

Cyrtodactylus huongsonensis sp. nov.

Holotype. Adult male (IEBR A.2011.3) collected on 13 June 2011 by Vinh Quang Luu, Hieu Van Pham, and Nghia Van Ha in Huong Son limestone forest (25o35’N, 105o45’E), My Duc District, Hanoi, northern Vietnam at an eleva- tion of ca. 120 m a.s.l. (Figs. 1–4). Paratype. Adult female (ZFMK 92293) the same collection data as for the holotype (Fig. 4). Diagnosis. A medium-sized Cyrtodactylus with a maximum SVL of 89.8 mm, distinguished from all conge- ners by the combination of the following characters: 1) dorsal pattern consisting of a dark nuchal loop, neck band and five in part irregularly shaped transverse body bands between limbs; 2) two enlarged lateral chinshields in con- tact with first postmental pair; 3) dorsal tubercles present on occiput, body, forearms, hind limbs and tail base; 4) 14–16 irregular dorsal tubercle rows; 5) ventrals in 41–48 longitudinal rows at midbody; 6) lateral skin folds present, lacking tubercles; 7) six precloacal pores plus in total 17 femoral pores in males separated by 8–12 pore- less scales; 8) enlarged femoral scales present; 9) enlarged precloacal scales present; 10) precloacal groove absent; 11) 3 postcloacal spurs in males; 12) subcaudal scales transversally enlarged. Description of holotype. Size medium (SVL 89.8 mm, TaL 78.0 mm, regenerated), distance from posterior corner of eye to anterior margin of ear including ciliaria 8.4 mm, maximum horizontal ear diameter 1.9 mm; for further measurements see Table 1. Rostral wider than high (RW 3.7 mm, RH 2.2 mm, RW/RH 1.2) with an inverse Y-shaped median suture; supralabials 10/12; infralabials 10/11; nares bordered by rostral anteriorly, first supralabial laterally and four nasals posteriorly; supranasals separated from each other by two nasorostrals and a rectangular internasal; medial snout scales granular, those in contact with and nearby supralabials flattened and larger than medial scales; upper anterior ciliaries 3–4 times larger than posterior cilaries; head scales somewhat granular, smaller than median snout scales; anterior supraocular region, back of head and temporal region with rounded, 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 labials, one pair of enlarged gular scales, with four small gular scales in between them; dorsal scales somewhat granular, about as large as medial snout scales; dorsal tubercles round, conical, surrounded by 9–10 granular scales, tubercles forming approximately 14 irregular longitudinal rows at midbody; ventral scales smooth, medial scales 2–4 times larger than dorsal gran- ules, 48 longitudinal rows at midbody; lateral folds present, without interspersed tubercles; upper arm lacking enlarged tubercles, lower arm with some in part indistinct conical tubercles; dorsal hind limb covered with scat-

40 · Zootaxa 3129 © 2011 Magnolia Press LUU ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. tered, smooth, flat to conical tubercles; femoral pores bearing scales distinctly enlarged, separated from precloacal, pore-bearing scales by 8/12 poreless femoral scales; 9/7 enlarged femoral scales, 10/7 femoral pores, six precloacal pores in an angular series; fingers and toes lacking distinct webbing; lamellae numbering 15/15 under first finger, 19/17 under fourth finger, 15/15 under first toe, and 20/20 under fourth toe; claws surrounded by a small scale on upper and a large scale on lower sides; precloacal region covered by a patch of approximately 29 enlarged scales below precloacal, pore-bearing scales; precloacal groove absent; 3/3 enlarged postcloacal tubercles on lateral sur- face of hemipenial swelling; dorsum of tail bearing distinct tubercles at base only, remainder of tail smooth, regen- erated; subcaudals distinctly enlarged, flat, smooth.

TABLE 1. Selected measurements and morphological characters of the male holotype and female paratype of Cyrtodactylus huongsonensis sp. nov.; measurements in mm, * = regenerated tail. IEBR A.2011.3 ZFMK 92293 SVL 89.8 73.4 TaL 78.0* 90.5 HH 10.1 9.0 HL 24.7 22.2 HW 17.2 15.1 OD 5.6 5.1 SE 10.2 9.9 SL 10/12 13/12 IL 10/11 12/12 N4/4 4/4 IN 1 1 CIL 26/26 28/27 PM 2 2 DTR 14 16 GST 8-10 8-10 V48 41 SLB 191 170 EFS 9/7 9/9 FP 10/7 8/7 PP 6 8 PAT 3/3 2/2 LD4 19/17 19/19 LT4 20/20 23/22

Coloration in ethanol. Ground colouration light brownish-grey, with dark brownish-black dorsal pattern; dor- sal head surface with irregular dark blotches, largest at occiput; dark canthal stripe, extending from nostril to eye; postocular streak only distinct on the right side, continuing to contact a somewhat irregularly shaped nuchal loop; postocular streak and nuchal loop bordered by thin light line; neck with a short dark transversal band; five more distinct dark transverse bands between limbs; dark transverse body bands somewhat irregularly shaped, with dark and light borders, fading towards sides; interspaces between dark dorsal bands with dark reticulation or blotches; dark tubercles in the nuchal loop and body bands, whereas light tubercles comprise the light lines and interspaces; flanks with dark blotches; upper surfaces of limbs with dark stripes and reticulations; dorsal surface of regenerated tail grey; gular region yellowish-cream; venter cream and lower tail surface light grey. For coloration in life see Fig. 1. The light lines bordering the nuchal loop and dark dorsal bands are yellow in life, as are the light dorsal tubercles; ground colouration in life is light greyish-brown.

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FIGURE 1. Portrait (A) and dorsal aspect (B) of the male holotype of Cyrtodactylus huongsonensis sp. nov. in life (IEBR A.2011.3). Photos V.Q. Luu

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FIGURE 2. Preserved holotype of Cyrtodactylus huongsonensis sp. nov. (IEBR A.2011.3): head in lateral (A), dorsal (B) and ventral (C) view, as well as mid-dorsum (D). Photos T. Ziegler.

Variation of paratype. For the variation in colouration of the female paratype see Fig. 4 and for meristics see Table 1. The internasal is very small and located between the anterior edges of the nasorostrals. The tail is original and has distinctly broadened subcaudals. The female paratype is somewhat smaller, but due to egg development we classify it as an adult, which is also indicated by the tail colouration. The dorsal surface of the original tail has 11 dark and 11 light bands, of which the last four dark bands are only faintly discernible. The disappearing light tail tip indicates the transition from juvenile to adult stage (see Ziegler et al. 2010). Concerning the colour pattern, the pos- tocular streaks are well developed on both sides of the female paratype. Here, also the lateral dark blotches are more distinct than it is the case in the male holotype. The dark blotches on the occiput are fused to a black v-shaped transverse line. The underside of the tail is dark grey, with the tip being light. With respect to sexual dimorphism, the female lacks hemipenial swellings at the tail base, the precloacal and femoral pores are only faintly discernible, and the postcloacal tubercles are less pronounced and fewer in number (2/2 versus 3/3 in the male holotype). 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 & Gris- mer 2010, Ngo & Pauwels 2010, Sumontha et al. 2010, Ziegler et al. 2010, David et al. 2011, Iskandar et al. 2011, Ngo 2011, Schneider et al. 2011). Cyrtodactylus huongsonensis sp. nov. differs from its Vietnamese congeners by the following characters (for details see Table 2): Cyrtodactylus huongsonensis sp. nov. has enlarged subcaudal scales and thus differs from the following species, which lack enlarged subcaudals: C. cattienensis Geissler, Nazarov, Orlov, Böhme, Phung, Nguyen & Ziegler, 2009, C. cryptus Heidrich, Rösler, Vu, Böhme & Ziegler, 2007, C. huynhi Ngo & Bauer, 2008, C. irregularis (Smith, 1921), C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, 2008, and C. ziegleri Nazarov, Orlov, Nguyen & Ho, 2008. Cyrtodactylus huongsonensis sp. nov. has enlarged femoral scales, which are lacking in the following species: C. badenensis Nguyen, Orlov & Darevsky, 2006, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, 2007, C. grismeri Ngo, 2008, and C. nigriocularis Nguyen, Orlov & Dare- vsky, 2006. Cyrtodactylus huongsonensis sp. nov. has femoral pores, which are absent in the following species:

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FIGURE 3. Undersides of right hand (A) and foot (B) of the preserved holotype of Cyrtodactylus huongsonensis sp. nov. (IEBR A.2011.3). Photos T. Ziegler.

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FIGURE 4. A: Cloacal region of the preserved holotype of Cyrtodactylus huongsonensis sp. nov. (IEBR A.2011.3); precloacal and femoral pores were marked with black. B: Dorsal view of preserved male holotype (top) and female paratype (ZFMK 92293, bottom) of Cyrtodactylus huongsonensis sp. nov. Photos T. Ziegler.

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C. eisenmanae Ngo, 2008, C. hontreensis Ngo, Grismer & Grismer, 2008, C. intermedius (Smith, 1917), C. martini Ngo, 2011, C. paradoxus (Darevsky & Szczerbak, 1997), and C. phuquocensis Ngo, Grismer & Grismer, 2010. Cyrtodactylus huongsonensis sp. nov. has 15–17 femoral pores in total and thus differs from C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 2007 (6), C. takouensis Ngo & Bauer, 2008 (0–2), and C. yangbayensis Ngo & Chan, 2010 (0–2). Cyrtodactylus huongsonensis sp. nov. has precloacal pores separated from femoral pores, which are a contiguous series in C. phongnhakebangensis Ziegler, Rösler, Herrmann & Vu, 2003, and C. roesleri Ziegler, Nazarov, Orlov, Nguyen, Vu, Dang, Dinh & Schmitz, 2010. Cyrtodactylus huongsonensis sp. nov. has three postcloacal tubercles in males and 41–48 ventrals and thus differs from C. bichnganae Ngo, 2010, that has only two postcloacal tubercles in males and 30–31 ventrals. Cyrtodactylus huongsonensis sp. nov. lacks tuber- cles on the lateral skin fold and on the dorsal tail surface and thus differs from C. condorensis (Smith, 1921); in addition, the latter species only has precloacal pores (4–7) and a blotched dorsal pattern. With respect to the remaining Cyrtodactylus, Cyrtodactylus huongsonensis sp. nov. has transversely enlarged subcaudals and thus differs from the following species which lack enlarged subcaudals: C. adleri Das, 1997, C. aequalis Bauer, 2003, C. agusanensis (Taylor, 1915), C. annulatus (Taylor, 1915), C. batucolus Grismer, Chan, Grismer, Wood & Belabut, 2008, C. biordinis Brown & McCoy, 1980, C. brevidactylus Bauer, 2002, C. buchardi David, Teynié & Ohler, 2004, C. cavernicolus Inger & King, 1961, C. derongo Brown & Parker, 1973, C. fumosus (Müller, 1895), C. gansi Bauer, 2003, C. halmahericus Mertens, 1929, C. irianjayaensis Rösler, 2000, C. jamban- gan Welton, Siler, Diesmos & Brown, C. jellesmae (Boulenger, 1897), C. lateralis (Werner, 1896), C. loriae (Bou- lenger, 1898), C. louisiadensis (De Vis, 1892), C. malayanus (De Rooij, 1915), C. mandalayensis Mahony, 2009, C. marmoratus Gray, 1831, C. matsuii Hikida, 1990, C. murua Kraus & Allison, 2006, C. novaeguineae (Schlegel, 1837), C. nuaulu Oliver, Edgar, Mumpuni, Iskandar & Lilley, 2009, C. pantiensis Grismer, Chan, Grismer, Wood & Belabut, 2008, C. papuensis (Brongersma, 1934), C. philippinicus (Steindachner, 1867), C. pubisulcus Inger, 1957, C. quadrivirgatus Taylor, 1962, C. sadleiri Wells & Wellington, 1985, C. semenanjungensis Grismer & Leong, 2005, C. seribuatensis Youmans & Grismer, 2006, C. sermowaiensis (de Rooij, 1915), C. serratus Kraus, 2007, C. stresemanni Rösler & Glaw, 2008, C. sworderi (Smith, 1925), C. tautbatorum Welton, Siler, Diesmos & Brown, 2009, C. tiomanensis Das & Lim, 2000, C. tuberculatus (Lucas & Frost, 1900), C. wakeorum Bauer 2003, C. wetariensis (Dunn, 1927), C. yoshii Hikida, 1990, C. zhaoermii Shi & Zhao, 2010, and C. zugi Oliver, Tjarhan, Mumpuni, Krey & Richards, 2008. Cyrtodactylus huongsonensis sp. nov. has femoral pores and thus differs from the following species which lack femoral pores: C. angularis (Smith, 1921), C. aurensis Grismer, 2005, C. ayeyarwadyensis Bauer, 2003, C. chan- homeae Bauer, Sumontha & Pauwels, 2003, C. chrysopylos Bauer, 2003, C. consobrinoides (Annandale, 1905), C. cracens Batuwita & Bahir, 2005, C. deveti (Brongersma, 1948), C. edwardtaylori Batuwita & Bahir, 2005, C. elok Dring, 1979, C. feae (Boulenger, 1893), C. fraenatus (Günther, 1864), C. ingeri Hikida, 1990, C. jarujini Ulber, 1993, C. khasiensis (Jerdon, 1870), C. malcolmsmithi (Constable, 1949), C. oldhami (Theobald, 1876), C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, 2011, C. papilionoides Ulber & Grossmann, 1991, C. peg- uensis (Boulenger, 1893), C. ramboda Batuwita & Bahir, 2005, C. rubidus (Blyth, 1860), C. soba Batuwita & Bahir, 2005, C. subsolanus Batuwita & Bahir, 2005, C. sumonthai Bauer, Pauwels & Chanhome, 2002, and C. var- iegatus (Blyth, 1859). Cyrtodactylus huongsonensis sp. nov. has six precloacal pores plus in total 17 femoral pores in males, which are separated by poreless scales and thus differs from the following species which have a contiguous series of pre- cloacal-femoral pores: C. epiroticus Kraus, 2008 (60–82), C. klugei Kraus, 2008 (66–76), C. lomyenensis Ngo & Pauwels, 2010 (32–40), C. robustus Kraus, 2008 (75–85), C. salomonensis Rösler, Richards & Günther, 2007 (71– 72), C. tamaiensis Mahony, 2009 (40), and C. tripartitus Kraus, 2008 (64–78). Cyrtodactylus huongsonensis sp. nov. has six precloacal pores in males and 8 precloacal pores in females and thus differs from the following species which have distinctly higher precloacal pore counts: C. annandalei Bauer, 2003 (11–12), C. capreoloides Rösler, Richards & Günther, 2007 (13), C. durio Grismer et al., 2010 (12), C. inter- digitalis Ulber, 1993 (14), C. russelli Bauer, 2003 (15), C. spinosus Linkem, McGuire, Hayden, Setiadi, Bickford & Brown (12–13), and C. teyniei David, Nguyen, Schneider & Ziegler, 2011 (14 in the single known specimen, an adult female). The following Cyrtodactylus species differ from Cyrtodactylus huongsonensis sp. nov. by the absence of pre- cloacal and femoral pores in both sexes: C. batik Iskandar, Rachmansah & Umilaela, 2011, C. darmandvillei (Weber, 1890), C. gordongekkoi (Das, 1993) (see Biswas 2007), C. jarakensis Grismer, Chan, Grismer, Wood &

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Belabut, 2008, C. laevigatus (Darevsky, 1964), C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome 2004, and C. wallacei Hayden, Brown, Gillespie, Setiadi, Linkem, Iskandar, Umilaela, Bickford, Riyanto, Mumpuni & McGuire, 2008. Cyrtodactylus huongsonensis sp. nov. has 41–48 ventral scales at midbody and thus differs from C. agamensis (Bleeker, 1860) (67), C. consobrinus (Peters, 1871) (65–70), C. erythrops Bauer et al., 2008 (28), C. gubernatoris (Annandale, 1913) (30), C. leegrismeri Chan & Norhayati, 2010 (27–35), C. macrotuberculatus Grismer & Norhayati, 2008 (19–22), C. pulchellus Gray, 1828 (33–35), C. slowinskii Bauer, 2002 (27–32), C. tigroides Bauer, Sumontha & Pauwels, 2003 (34), and C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler, 2010 (31–35). Cyrtodactylus huongsonensis sp. nov. has dorsal tail surface without tubercles and thus differs from C. aaroni Günther & Rösler, 2003, C. brevipalmatus (Smith, 1923), C. redimiculus King, 1962, C. baluensis (Mocquard, 1890), and C. mimikanus (Boulenger, 1914). Cyrtodactylus huongsonensis sp. nov. has 14–16 dorsal tubercle rows and thus differs from C. auribalteatus Sumontha, Panitvong & Deein, 2010 (22–24), and C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, 2010 (18–22). In addition, Cyrtodactylus huongsonensis sp. n. differs from the representatives of the subgenus Geckoella by the presence of transversally enlarged subcaudals and by having precloacal pores in both sexes: C. (Geckoella) albofasciatus (Boulenger, 1885), C. (Geckoella) collegalensis (Beddome, 1870), C. (Geckoella) deccanensis (Günther, 1864), C. (Geckoella) jeyporensis (Beddome, 1877), C. (Geckoella) nebulosus (Beddome, 1870), C. (Geckoella) triedrus (Günther, 1864), and C. (Geckoella) yakhuna (Deraniyagala, 1945). Distribution. Cyrtodactylus huongsonensis sp. nov. is currently known only from the type locality in Vietnam (Fig. 5). Etymology. We name this species after its type locality, Huong Son limestone forest, which is a tourist site and not yet protected. Huong Son tourist site covers an area of a thousand hectares and includes a complex of moun- tains, rivers and streams, villages, pagodas, and grottoes surrounded by the Huong Tich Mountain Range, north of the Truong Son Range. Ecological notes. The type specimens were found at night on karst outcrops in limestone forest, at an elevation of ca. 120 m a.s.l. The left testis of the male holotype measured 8.0 mm in length. The somewhat smaller female paratype contained further developed eggs of up to 3.4 mm diameter.

Discussion

The phylogenetic relationships of Cyrtodactylus huongsonensis sp. nov. with its congeners are not yet known, as is the phylogeny of the entire genus, which consists of more than 140 species. Most recently, C. bichnganae was described by Ngo & Grismer (2010) from the karst forests of Son La Province, which is westward from Hanoi, with Phu Tho and Hoa Binh provinces in between. Phenetically, this geographically close species resembles Cyrto- dactylus huongsonensis sp. nov., but there are a number of morphologically distinguishing characters between these karst adapted species, such as two enlarged lateral chinshields in contact with first postmental pair in Cyrto- dactylus huongsonensis sp. nov. versus four in C. bichnganae, 41–48 versus 30–31 ventrals, 10 versus six precloa- cal pores in males, and three postcloacal tubercles in males versus two. The distinctness of both taxa is also discernible by the width of the diastema between precloacal and femoral pores: these pores are separated by 8–12 poreless scales on each side in Cyrtodactylus huongsonensis sp. nov., whereas precloacal and femoral pores are separated by only two poreless scales on each side in C. bichnganae. There are also differences in the dorsal colour pattern, such as a clearer banded pattern in C. bichnganae, without distinct prominent dark interspaces as it is the case in Cyrtodactylus huongsonensis sp. nov. Also the light dorsal tubercles are more prominent and conspicuous in the new species. The type locality of Cyrtodactylus huongsonensis sp. nov. in Hanoi is about 150 km distant from the type locality of C. bichnganae in Son La Province. Both karst formations (Hanoi versus Son La) are loca- ted in the western side of the Red River but they are separated from each other by the Da River. Whatsoever, only future field work will help to identify proper species distribution boundaries and molecular sampling is certainly required to both understand the phylogenetic relations of Cyrtodactylus huongsonensis sp. nov. and create a relia- ble phylogeny of Cyrtodactylus.

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FIGURE 5. Map showing the type locality of Cyrtodactylus huongsonensis sp. nov. in Vietnam.

Acknowledgements

We thank C.X. Le (IEBR, Hanoi) for the loan of specimens and the kind permission to donate a type specimen to the herpetological collection of the ZFMK, Bonn, Germany. We also would like to thank A. Gawor (Cologne Uni- versity / Cologne Zoo) for the transfer of specimens and the kind arrangement of the colour plates. Thanks also to L.L. Grismer (La Sierra University, Riverside) and O.S.G. Pauwels (Institut Royal des Sciences naturelles de Belgique, Brussels) for reviewing the manuscript.

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