Cyrtodactylus kunyai (Squamata: Gekkonidae), a new ...

Zootaxa 3821 (2): 253–264 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3821.2.5 http://zoobank.org/urn:lsid:zoobank.org:pub:803D24E1-2FA8-431E-B9D4-D20A0D53EBB0

Cyrtodactylus kunyai (Squamata: Gekkonidae), a new cave-dwelling Bent-toed Gecko from Loei Province, northeastern Thailand OLIVIER S. G. PAUWELS1,5, MONTRI SUMONTHA2, KAWEESAK KEERATIKIAT3 & EAKARIT PHANAMPHON4 1

Département des Vertébrés Récents, Institut Royal des Sciences naturelles de Belgique, Rue Vautier 29, B-1000 Brussels, Belgium. E-mail: [email protected] 2 Ranong Marine Fisheries Station, 157 Saphanpla Rd., Paknam, Muang, Ranong 85000, Thailand. E-mail: [email protected] 3 264/45-46 Suksawat 15, Bangpakok, Rajburana, Bangkok 10140, Thailand 4 300 Ratchasima Rd., Dusit, Bangkok 10300, Thailand 5 Corresponding author

Abstract We describe a new cave-dwelling Cyrtodactylus from Suan Hin Pha Ngam, Nong Hin District, Loei Province, northeastern Thailand, characterized by a maximal known snout-vent length of 87.9 mm, a banded dorsal pattern with a medially interrupted nuchal loop and four or five brown bands between nuchal loop and hind limb insertions and three bicolored band interspaces between limbs insertions, a dark orangeish iris, a continuous series of enlarged femoro-precloacal scales with 5–6 femoral pores on each side separated by a diastema from 3 precloacal pores in males (no pores in females), 19 irregularly arranged dorsal longitudinal tubercle rows at midbody, 34 ventral scale rows between ventrolateral skin folds, transversely enlarged subcaudal plates, and no precloacal groove. Key words: Cyrtodactylus kunyai sp. nov., taxonomy, limestone cave, Little Kunming

Introduction The limestone hills of Suan Hin Pha Ngam (literally translating as the ‘‘Stone Garden of the Beautiful Cliff’’) are a famous tourist attraction in Loei Province. They were nicknamed ‘’Little Kunming’’ because they remind of the dramatic landscapes found in the southern Chinese Yunnan Province. In the course of fieldwork related to a systematic and phytogeographic review of the Impatiens (Balsaminaceae) of Thailand, two of us (KK and EP) found a population of Cyrtodactylus in a cave of Suan Hin Pha Ngam. They differ from all known species by a combination of color pattern and scalation features, and are consequently described here as a new species.

Material and methods Measurements and meristic counts follow Sumontha et al. (2012) and Pauwels et al. (2013). Paired meristic characters are given left/right. Numbers of supralabial and infralabial scales are counted from the largest scale immediately posterior to the dorsal inflection of the posterior portion of the upper jaw to the rostral and mental scales, respectively. The number of longitudinal rows of body tubercles was counted transversely across the center of the dorsum from one ventrolateral skin fold to the other. The number of longitudinal rows of ventral scales was counted transversely across the center of the abdomen from one ventrolateral skin fold to the other. The numbers of subdigital lamellae beneath the toes were counted from the base of the first phalanx to the claw. The following measurements were taken with a digital caliper to the nearest 0.1 mm: AG: axilla to groin length, taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its Accepted by A. Bauer: 28 May 2014; published: 20 Jun. 2014

253

insertion point on the body; EarL: ear length, the greatest horizontal distance of the ear opening; ForeaL: forearm length, taken on the dorsal surface from the posterior margin of the elbow while flexed 90° to the inflection of the flexed wrist; HeadH: head height, the maximum depth of head from the occiput to the throat; HeadL: head length, from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; HeadW: head width, measured at the angle of the jaws; Internar: internarial distance, measured between the nares across the rostrum; Interorb: interorbital distance, measured between the anterior edges of the orbits; NosOrb: nostril to orbit distance, from the posterior margin of the external nares to the anterior margin of the orbit; OrbD: orbit diameter, the greatest horizontal diameter of the orbit; OrbEar: orbit to ear distance, from the anterior edge of the ear opening to the posterior edge of the orbit; SnEye: snout to eye distance, from the tip of the snout to the anteriormost margin of the orbit; SVL: snout-vent length, taken from the tip of snout to the vent; TailL: tail length, taken from the vent to the tip of the tail, original or regenerated; TailW: tail width, taken at the base of the tail immediately posterior to the postcloacal swelling; TibiaL: tibia length, taken on the ventral surface from the posterior surface of the knee while flexed 90° to the base of heel. Meristic characters abbreviations: DorTub: dorsal tubercles; FemP: femoral pores; IL: infralabial scales; InterorbSc: interorbital scales; PreclP: precloacal pores; SL: supralabial scales; Ven: ventral scales. Museum and other acronyms: IRSNB: Institut Royal des Sciences naturelles de Belgique, Brussels; MS: Montri Sumontha’s field number series; THNHM: Thailand Natural History Museum, National Science Museum, Technopolis, Pathum Thani.

Results Cyrtodactylus kunyai sp. nov. (Figs 1–8) Holotype. THNHM 2560 (field number MS 270); adult male from a limestone cave in Suan Hin Pha Ngam (17°03.062’N, 101°44.588’E), Amphoe (= District) Nong Hin, Loei Province, northeastern Thailand. Collected by Kaweesak Keeratikiat on 23 June 2008. Diagnosis. Cyrtodactylus kunyai sp. nov. can be distinguished from all other congeneric species by its maximal known SVL of 87.9 mm; 19 longitudinal rows of dorsal tubercles; 34 ventral scale rows between ventrolateral skin folds; a continuous series of 39 enlarged femoro-precloacal scales, including 5–6 pore-bearing scales on each femur separated by a diastema from 3 pore-bearing precloacal scales (no pores in females); no precloacal groove nor depression; transversely enlarged subcaudal scales; a dark orangeish iris; adults with four or five brown bands between nuchal loop and hind limb insertions and 3 bi-colored band interspaces between limb insertions and a medially interrupted nuchal loop. Description of holotype. Adult male. SVL 87.9 mm. Tail original, TailL 107.2 mm. Head relatively long (HeadL 24.8 mm; HeadL/SVL ratio 0.28), wide (HeadW 16.1 mm; HeadW/HeadL ratio 0.65), not markedly depressed (HeadH 10.1 mm; HeadH/HeadL ratio 0.41, distinct from slender neck. Loreal region weakly inflated, canthus rostralis not prominent. Snout elongate (SnOrb 9.8 mm; SnOrb/HeadL ratio 0.40), rounded, longer than orbit diameter (OrbD 6.6 mm; OrbD/SnOrb ratio 0.67); scales on snout small, rounded to oval, granular to weakly conical, mostly homogeneous, larger than those on crown, interorbital and occipital regions. Eye large (OrbD/ HeadL ratio 0.27); pupil vertical with crenelated margins; supraciliaries short (anteriormost longest), those at posterior part of orbit bearing small conical spines. Ear opening vertically elliptical, small (EarL 1.1 mm; EarL/ HeadL ratio 0.04); orbit to ear distance subequal to orbit diameter (OrbEar 6.5 mm; OrbEar/OrbD ratio 0.98; NosOrb 7.5 mm; Internar 2.6 mm; Interorb 4.7 mm). Rostral wider (3.5 mm) than deep (2.4 mm), rostral crease about a third of rostral height. Two enlarged supranasals in broad contact with one another. Rostral in contact with first supralabials, nostrils, and supranasals. Nostrils oval, dorsolaterally directed, each surrounded by supranasal, rostral, first supralabial and two enlarged postnasals. Three or four rows of small scales separate orbit from supralabials. Mental triangular, wider (3.4 mm) than deep (2.3 mm). A single pair of greatly enlarged postmentals in broad contact behind mental, each postmental bordered anteromedially by mental, anterolaterally by first infralabial, posterolaterally by two enlarged lateral chinshields, and posteriorly by four granules, the medialmost of which also contacting the other postmental. Supralabials to mid-orbital position 10/9, enlarged supralabials to angle of jaws 13/12. Infralabials 10/10. Interorbital scale rows across narrowest point of frontal bone 27. Gular region with homogeneous, smooth, juxtaposed granular scales.

254 · Zootaxa 3821 (2) © 2014 Magnolia Press

PAUWELS ET AL.

FIGURE 1. Live adult male holotype of Cyrtodactylus kunyai sp. nov., general dorsal view. Photo. taken ex situ by M. Sumontha.

FIGURE 2. Live adult male holotype of Cyrtodactylus kunyai sp. nov., head view. Photo. taken ex situ by M. Sumontha.

A NEW CAVE GECKO FROM EASTERN THAILAND

Zootaxa 3821 (2) © 2014 Magnolia Press ·

255

FIGURE 3. Live adult male holotype of Cyrtodactylus kunyai sp. nov., ventral view. Photo. taken ex situ by M. Sumontha.

FIGURE 4. Live subadult male Cyrtodactylus kunyai sp. nov. Photo. taken in situ by K. Keeratikiat; individual not collected.


PAUWELS ET AL.

FIGURE 5. Live adult female Cyrtodactylus kunyai sp. nov. Photo. taken ex situ by M. Sumontha; individual not collected.

FIGURE 6. Live adult female Cyrtodactylus kunyai sp. nov., ventral view, showing the absence of femoral or precloacal pores or pits. Photo. taken ex situ by M. Sumontha; individual not collected.



257

FIGURE 7. Another live adult female Cyrtodactylus kunyai sp. nov. Photo. taken in situ by K. Keeratikiat; individual not collected.

FIGURE 8. Live juvenile Cyrtodactylus kunyai sp. nov. Photo. taken in situ by K. Keeratikiat; individual not collected.


PAUWELS ET AL.

FIGURE 9. Geographical location of the type locality of Cyrtodactylus kunyai sp. nov. (black dot) in Nong Hin District, Loei Province; map by W. Sodoab.

Body slender, moderately elongate (AG 39.7 mm; AG/SVL ratio 0.45) with relatively well-defined, nondenticulate ventrolateral folds. Dorsal scales weakly heterogeneous, flat or slightly domed; irregularly distributed tubercles (about three times size of adjacent scales) extending from shoulder region onto tail base, smaller tubercles on postocular region, crown, occiput and nape; some tubercles bearing a low keel, those on flanks flat, most lacking a keel; tubercles in 19 irregular rows at midbody, typically separated from one another by two dorsal granules. Ventral scales larger than dorsals, smooth, oval and subimbricate, larger in precloacal region. Midbody scale rows across belly between ventrolateral folds 34. A continuous row of 39 enlarged femoro-precloacal scales, as follows, from left to right: 5 pore-bearing femoral scales + a diastema of 12 unpitted poreless scales + 3 porebearing preanal scales + a diastema of 12 unpitted poreless scales 6 pore-bearing femoral scales + 1 unpitted



259

poreless femoral scale. Distal femoral scales about three times the size of adjacent anterior femoral scales; proximal femoral scales of the femoro-precloacal series smaller, about the size of adjacent anterior femoral scales. No precloacal groove nor depression. Postcloacal spurs each bearing two enlarged, conical scales, the anterior one largest. Scales on palm and sole smooth, rounded to oval or hexagonal, slightly domed. Scalation on dorsal surface of hind limbs similar to body dorsum, with enlarged conical tubercles interspersed among smaller scales; no enlarged tubercles on fore limbs. Fore and hind limbs relatively long, slender (ForeaL 14.8 mm, ForeaL/SVL ratio 0.17; TibiaL 17.9 mm, TibiaL/SVL 0.20). Digits long, slender, inflected at interphalangeal joints, all bearing robust, slightly recurved claws. Basal subdigital lamellae broad, ovoid to rectangular, without scansorial surfaces (5-7-7-69 right manus, 5-7-8-9-9 right pes); narrow lamellae distal to digital inflection and not including ventral claw sheath: 9-10-9-11-10 (right manus), 10-12-12-11-11 (right pes); weak interdigital webbing between digits and toes (except between toes IV and V). Tail original, gently tapering to pointed tip, longer than SVL (TailL/SVL ratio 1.22); TailW 6.0 mm. Median subcaudal plates enlarged.

FIGURE 10. Type locality and macro-habitat of Cyrtodactylus kunyai sp. nov. Photo by K. Keeratikiat.

Coloration in life. Dorsal ground color of head, dorsum, limbs and tail beige. Upper surface of head with a brown reticulum. One bi-colored interspace on neck between brown nuchal loop and a brown band above anterior insertion of forelimbs, dark brown anteriorly and posteriorly, beige with irregular transversely arranged dark brown spots in their middle. Another such band interspace above shoulders, three between limbs insertions, one above hind legs insertion, four on anterior half of tail; between limbs insertions these band interspaces forked at their base (Figs 1–2). On dorsum bands of the same length as band interspaces. Nuchal loop connecting the orbits interrupted medially by a yellowish stripe connecting the neck band interspace to the reticulum on upper surface of head. Upper surfaces of fore and hind limbs with irregular and discontinuous transversal bands and bi-colored band interspaces. Posterior part of tail grayish mottled with beige and dark brown. Iris dark orangeish. Throat, ventrum


PAUWELS ET AL.

and underside of fore limbs uniformly whitish, underside of hind limbs darker, except the series of enlarged femoro-precloacal scales, whitish; underside of tail brown (Fig. 3). Variation. Besides the holotype, four other individuals (not preserved) were photographed and examined: a subadult male (Fig. 4), two adult females (Figs 5–7) and a juvenile (Fig. 8). The adults and subadult show a pattern similar to the holotype, with three bicolored band interspaces between limbs insertions, a dark orangeish iris and a medially interrupted nuchal loop. The brown band between the nuchal loop and the brown band above shoulders is variably present (Figs 1–2, 4–8). The juvenile shows a white posterior half of tail, similarly to the subadult male; it also shows a partly interrupted nuchal loop, even if it is less obvious than in adults. The females have a continuous series of enlarged femoro-precloacal scales but lack femoral or precloacal pores or pits (Fig. 6). Distribution and natural history. The species is currently known only from its type locality where it is common. It was found inside small limestone caves and crevices and on nearby outcrops. It is nocturnal. Outside the caves it was found in syntopy with Cyrtodactylus interdigitalis Ulber, Dixonius siamensis (Boulenger), Gehyra fehlmanni (Taylor) and Gekko gecko (Linnaeus) (Gekkonidae). Etymology. The specific epithet, formed in the masculine genitive, honors our friend and colleague Kirati Kunya, curator of the reptile collection of the Nakhonratchasima Zoo, Khorat, for his contributions to the herpetology of Thailand. We suggest the following common names: Took-kai Kirati (Thai), Kunya’s bent-toed gecko (English) and Cyrtodactyle de Kunya (French).

Discussion We compared Cyrtodactylus kunyai sp. nov. with all species known from Cambodia, China, Laos, Myanmar, Thailand and Vietnam (see latest descriptions and intraspecific comparisons in Ulber & Grossmann 1991; Bauer 2002, 2003; Grismer 2008; Mahony 2009; Bauer et al. 2010; Ngo & Chan 2010; Ngo & Pauwels 2010; Nguyen et al. 2010; Chan-ard & Makchai 2011; David et al. 2011; Grismer et al. 2012; Ngo & Grismer 2012; Sumontha et al. 2010; 2012; Nguyen et al. 2013; Pauwels et al. 2013, 2014; Ziegler et al. 2013; Luu et al. 2014; Pauwels & Sumontha 2014; Phung et al. 2014; Schneider et al. 2014). Among the species from these six countries, the phylogenetic relationships of Cyrtodactylus kunyai sp. nov. are likely closer to the other species sharing the possession of precloacal and femoral pores separated by a diastema. From these latter species, Cyrtodactylus kunyai sp. nov. can be readily distinguished by a combination of the following characters: enlarged subcaudals (absent in C. aequalis Bauer, C. huynhi Ngo & Bauer, and C. ziegleri Nazarov, Orlov, Nguyen & Ho); 3 precloacal pores (9 in C. aequalis (female holotype), 11–12 in C. annandalei Bauer, six in C. auribalteatus Sumontha, Panitvong & Deein, ten in C. bichnganae Ngo & Grismer, 7–9 in C. brevipalmatus Smith, nine in C. caovansungi Orlov, Nguyen, Nazarov, Ananjeva & Nguyen, 9–11 in C. consobrinus (Peters), five or six in C. dumnuii Bauer, Kunya, Sumontha, Niyomwan, Pauwels, Chanhome & Kunya, six in C. huongsonensis Luu, Nguyen, Do & Ziegler, 7–9 in C. huynhi and C. kingsadai, 15 in C. russelli Bauer, nine in C. slowinskii Bauer, zero or one in C. thuongae Phung, Van Schingen, Ziegler & Nguyen, 2014, eight or nine in C. tigroides Bauer, Sumontha & Pauwels, 2003, 6-8 in C. yangbayensis Ngo & Chan, 5–8 in C. ziegleri); 19 longitudinal tubercle rows (24 in C. aequalis, 16–18 in C. annandalei, 22–24 in C. auribalteatus, 16–18 in C. bichnganae, 18 in C. brevipalmatus, 16–18 in C. caovansungi, 14–16 in C. huongsonensis, nine or ten in C. huynhi, 22 in C. russelli, nine or ten in C. takouensis, 16–18 in C. thuongae, 13 in C. tigroides, 20–23 in C. yangbayensis, 20–24 in C. ziegleri); a reticulated pattern on head, a medially interrupted nuchal loop, four or five brown bands between nuchal loop and hind limb insertions and three wide bi-colored inter-band spaces between limb insertions (complete nuchal loop, paired dark blotches on dorsum in C. aequalis; no reticulum on head, complete nuchal loop and unicolor band interspaces in C. annandalei; complete nuchal loop in C. auribalteatus; six or seven dark bands between nuchal loop and hind limb insertions in C. bichnganae; blotched dorsal pattern in C. brevipalmatus; two to four very thin, unicolored light band interspaces between limb insertions in C. consobrinus; five or six light band interspaces between limb insertions in C. dumnuii; complete nuchal loop, four light band interspaces between limb insertions in C. huongsonensis; complete nuchal loop, five or six irregular light band interspaces between limb insertions in C. huynhi; complete nuchal loop, four or five light band interspaces between limb insertions in C. kingsadai; irregular banded and blotched pattern in C. russelli; blotched dorsal pattern in C. slowinskii; complete nuchal loop, three or four unicolor yellow band interspaces between limb insertions in C. takouensis; blotched dorsal pattern in C.



261

thuongae; complete nuchal loop, four bi-colored band interspaces between limb insertions in C. tigroides; five or six irregular dark bands between limb insertions in C. yangbayensis; irregular banded-blotched dorsal pattern in C. ziegleri). In addition, the number of ventral scales of Cyrtodactylus kunyai sp. nov. (34) does not overlap with those of C. aequalis (24), C. annandalei (43), C. auribalteatus (38–40), C. bichnganae (30–31), C. brevipalmatus (35–44), C. caovansungi (38–44), C. consobrinus (58–65), C. dumnuii (40), C. huongsonensis (41–48), C. huynhi (43–46), C. kingsadai (39–46), C. russelli (35–41), C. slowinskii (27–32), C. takouensis (39–40) and C. yangbayensis (39–46). From C. khelangensis Pauwels, Sumontha, Panitvong & Varaguttanonda, 2014, which seems to be the closest species in terms of pattern and scalation, C. kunyai sp. nov. is distinguished by the absence of precloacal pores and femoral pits in females (Fig. 6) vs. their presence in C. khelangensis, its marbled caudal bands of the same length as the interspaces in the original tail (vs. immaculate bands two or three times longer than band interspaces in C. khelangensis), its longer dorsum band interspaces relative to the dorsal bands length, its light brown dorsal bands (vs. dark brown in C. khelangensis), the presence of longitudinally elongated dark marks within the band interspaces (absent in C. khelangensis) and a slightly smaller maximal known SVL (87.9 mm, vs. 95.3 mm for C. khelangensis) (see Pauwels et al. 2014). The pace at which cave and karst-dwelling Cyrtodactylus are being discovered and described from Thailand does not seem to decrease, at the contrary (Pauwels & Sumontha 2014). Pursuing the inventory of the Thai cave and karst Cyrtodactylus and analyzing and mitigating their conservation threats has become a conservation priority. Most of these species are micro-endemics, facing a set of specific conservation threats (Ellis & Pauwels 2012; Panitvong et al. 2012), one of them being intensive, uncontrolled tourism – the main potential issue that might threat Cyrtodactylus kunyai sp. nov. for instance. The type locality is indeed a popular touristic site, with an easy access through paths to caves and limestone cliffs. A further development of touristic infrastructure or increase of visitor numbers would probably represent a conservation threat to the species. As far as we know it has not been offered so far in the pet trade.

Acknowledgements We are grateful to Tanya Chan-ard (THNHM, Pathum Thani), Georges Lenglet and Sébastien Bruaux (IRSNB, Brussels) for providing access to the herpetological collections of their respective institutions, and to Watchira Sodoab for preparing the map.

References Bauer, A.M. (2002) Two new species of Cyrtodactylus (Squamata: Gekkonidae) from Myanmar. Proceedings of the California Academy of Sciences, 53 (7), 73–86. Bauer, A.M. (2003) Descriptions of seven new Cyrtodactylus (Squamata: Gekkonidae) with a key to the species of Myanmar (Burma). Proceedings of the California Academy of Sciences, 54 (25), 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 (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. 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 (1), 7–15. David, P., Nguyen, Q.T., Schneider, N. & Ziegler, T. (2011) A new species of the genus Cyrtodactylus Gray, 1827 from central Laos (Squamata: Gekkonidae). Zootaxa, 2833, 29–40. Ellis, M. & Pauwels, O.S.G. (2012) The bent-toed geckos (Cyrtodactylus) of the caves and karst of Thailand. Cave and Karst Science, 39 (1), 16–22. Grismer, L.L. (2008) On the distribution and identification of Cyrtodactylus brevipalmatus Smith, 1923 and Cyrtodactylus elok


PAUWELS ET AL.

Dring, 1979. The Raffles Bulletin of Zoology, 56(1), 177–179. 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. 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 Mahony, S. (2009) Taxonomic status of Cyrtodactylus khasiensis tamaiensis (Smith, 1940) and description of a new species allied to C. chrysopylos Bauer, 2003 from Myanmar (Reptilia: Gekkonidae). Hamadryad, 34 (1), 62–74. Ngo, V.T. & Chan, K.O. (2010) A new species of Cyrtodactylus Gray, 1826 (Squamata: Gekkonidae) from Khanh Hoa Province, Southern Vietnam. Zootaxa, 2504, 47–60. Ngo, V.T. & Grismer, L.L. (2012) A new endemic species of Cyrtodactylus Gray (Squamata: Gekkonidae) from Tho Chu Island, southwestern Vietnam. Zootaxa, 3228, 48–60. Ngo, V.T. & Pauwels, O.S.G. (2010) A new cave-dwelling species of Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Khammouane Province, southern Laos. Zootaxa, 2730, 44–56. Nguyen, S.N., Le, T.-N.T., Tran, T.A.D., Orlov, N.L., Lathrop, A., Macculloch, R.D., Le, T.-D.T., Jin, J.-Q., Nguyen, L.T., Nguyen, T.T., Hoang, D.D., Che, J., Murphy, R.W. & Zhan, 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, 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. Panitvong, N., Lauhachinda, V., Saithong, S. & Temchai, T. (2012) Ecology of Cyrtodactylus sumonthai Bauer, Pauwels, & Chanhome, 2002 (Reptilia: Squamata: Gekkonidae): a karst dwelling bent-toed gecko from south-eastern Thailand. The Raffles Bulletin of Zoology, 60 (2), 569–582. 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, 771, 1–11. Pauwels, O.S.G. & Chan-ard, T. (2006) Reptiles of Kaeng Krachan National Park, Western Thailand. Natural History Bulletin of the Siam Society, 54 (1), 89–108. Pauwels, O.S.G., Laohawat, O.-A., David, P., Bour, R., Dangsee, P., Puangjit, C. & Chimsunchart, C. (2000) Herpetological investigations in Phang-Nga Province, southern peninsular Thailand, with a list of reptile species and notes on their biology. Dumerilia, 4 (2), 123–154. 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 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 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 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 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. Ulber, T. & Grossmann, W. (1991) Ein weiterer neuer Gecko aus Zentral-Thailand: Cyrtodactylus papilionoides sp. nov. (Reptilia: Sauria: Gekkonidae). Sauria, 13 (1), 13–22. 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



263

APPENDIX. Comparative material examined. Cyrtodactylus astrum: see Grismer et al. (2012); C. auribalteatus: see Sumontha et al. (2010); C. brevipalmatus: see Pauwels & Chan-ard (2006); C. chanhomeae: see Bauer et al. (2003); C. dumnuii: see Bauer et al. (2010); C. erythrops: see Bauer et al. (2009); C. intermedius: IRSNB 17011, Nakhon Ratchasima, Thailand; C. khelangensis: see Pauwels et al. (2014); C. lekaguli: see Grismer et al. (2012); C. oldhami complex: see Pauwels & Chan-ard (2006) and Pauwels et al. (2000); C. peguensis: see Pauwels et al. (2000); C. phuketensis: see Sumontha et al. (2012); C. samroiyot: see Pauwels & Sumontha (2014); C. sanook: see Pauwels et al. (2013); C. sumonthai: see Bauer et al. (2002); C. thirakhupti: see Pauwels et al. (2004); C. tigroides: see Bauer et al. (2003).


PAUWELS ET AL.