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A new Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus) from Phetchaburi Province, Thailand

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A new Bent-toed Gecko, Cyrtodactylus phetchaburiensis sp. nov. is described from the Tha Yang District of Phetchaburi Province, western Thailand. It is a medium-sized Cyrtodactylus (SVL to at least 63.2 mm), with small, mostly keeled tubercles in 20 regular longitudinal rows on dorsum; 33 scales across mid-venter between lowest rows of flank tubercles; enlarged row of femoral scales present; five precloacal pores in male, femoral pores and precloacal groove absent; 5–6 broad basal lamellae and 11 narrow distal lamellae beneath digit IV of pes; and a single median row of transversely enlarged subcaudal scales present. It has a dorsal colour pattern of large, dark, diffusely-edged markings on a fawn background and a pair of dark scapular patches. The species is a member of the Central Indochinese (Thai-Myanmar) clade of Cyrtodactylus and is most closely related to C. oldhami (Theobald), from which it differs in colour pattern.
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Accepted by S. Carranza: 2 Feb. 2016; published: 10 Mar. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4088 (3): 409
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http://www.mapress.com/j/zt/
Article
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http://doi.org/10.11646/zootaxa.4088.3.6
http://zoobank.org/urn:lsid:zoobank.org:pub:B87BB845-FB3D-4536-8465-9DAB4B62AC53
A new Bent-toed Gecko (Squamata: Gekkonidae: Cyrtodactylus)
from Phetchaburi Province, Thailand
OLIVIER S. G. PAUWELS
1
, MONTRI SUMONTHA
2
& AARON M. BAUER
3,4
1
Département des Vertébrés Récents, Institut Royal des Sciences naturelles de Belgique, Rue Vautier 29, B-1000 Brussels, Belgium.
E-mail: osgpauwels@yahoo.fr
2
Ranong Marine Fisheries Station, 157 Saphanpla Rd., Paknam, Muang, Ranong 85000, Thailand.
E-mail: montri.sumontha@gmail.com
3
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA
4
Corresponding author. E-mail: aaron.bauer@villanova.edu
Abstract
A new Bent-toed Gecko, Cyrtodactylus phetchaburiensis sp. nov. is described from the Tha Yang District of Phetchaburi
Province, western Thailand. It is a medium-sized Cyrtodactylus (SVL to at least 63.2 mm), with small, mostly keeled tu-
bercles in 20 regular longitudinal rows on dorsum; 33 scales across mid-venter between lowest rows of flank tubercles;
enlarged row of femoral scales present; five precloacal pores in male, femoral pores and precloacal groove absent; 5–6
broad basal lamellae and 11 narrow distal lamellae beneath digit IV of pes; and a single median row of transversely en-
larged subcaudal scales present. It has a dorsal colour pattern of large, dark, diffusely-edged markings on a fawn back-
ground and a pair of dark scapular patches. The species is a member of the Central Indochinese (Thai-Myanmar) clade of
Cyrtodactylus and is most closely related to C. oldhami (Theobald), from which it differs in colour pattern.
Key words: Gekkonidae, Cyrtodactylus phetchaburiensis sp. nov., Cyrtodactylus oldhami, description, taxonomy, Thai-
land
Introduction
The Bent-toed Geckos (Cyrtodactylus Gray) constitute the most species rich reptile genus in tropical Asia, with
over 200 described species (Nazarov et al. 2014; Uetz 2015). The majority of these have been described in the last
decade, and many are limited to particular substrate types, such as caves and karst landscapes (e.g., Bauer et al.
2002, 2009, 2010; Ngo 2008; Sumontha et al. 2010; Schneider et al. 2011; Ellis & Pauwels 2012; Pauwels et al.
2014a,b; Grismer et al. 2014a, 2015). This has resulted in widespread microendemism and suggests that continued
herpetological exploration throughout the range of the genus, but especially in south-east Asia, will continue to
reveal additional species. In addition, widespread taxa of Cyrtodactylus, when investigated in detail, are frequently
revealed to be species complexes (e.g., Grismer et al. 2012, 2014b; Johnson et al. 2012; Nazarov et al. 2008, 2012;
Nguyen et al. 2013). Together, these trends have accounted for the explosion of Cyrtodactylus descriptions in
recent years.
Thirty-two Cyrtodactylus have been recorded from Thailand, fully half of which have been described since
2010. These recent descriptions have come from across the country and have included taxa in several different
clades (Wood et al. 2012). One of the lineages represented in Thailand is a group of taxa that occurs along the Thai-
Myanmar border. Molecular data have confirmed only three members of this clade, C. tigroides Bauer, Sumontha
& Pauwels, C. peguensis (Boulenger), and C. oldhami (Theobald), but additional taxa are expected, both because
previous sampling has been inadequate and because two of the constituent taxa, C. peguensis and C. oldhami, as
presently construed, have broad distributions and exhibit significant morphological variation, suggesting that they
may represent species complexes. Ulber (1993) illustrated and described specimens of C. oldhami from Kaeng
Krachan National Park, Phetchaburi Province that were atypical in colour pattern and some scale counts.
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Specimens from the park were also noted by Manthey & Grossmann (1997) and Nabhitabhata et al. (2004) and
were subsequently referred to as C. cf. oldhami (Pauwels & Chan-ard 2006; Pauwels et al. 2009). Subsequent field
work in Phetchaburi Province has revealed a new species of Cyrtodactylus that appears to be allied to C. oldhami
and to the Kaeng Krachan geckos and which we here describe.
Materials and methods
The following measurements were taken with digital calipers to the nearest 0.1 mm following the methods of Bauer
(2002, 2003): CrusL: crus length; EarL: ear length; EyeEar: eye to ear distance; ForeaL: forearm length; HeadH:
head height; HeadL: head length; HeadW: head width; Internar: internarial distance; Interorb: interorbital distance
measured between supraciliary scale rows; NarEye: nares to eye distance; OrbD: orbital diameter; SnEye: snout to
eye distance; SVL: snout-vent length; TailL: tail length; TailW: tail width; TrunkL: trunk length. Basal subdigital
lamellae were counted from the most proximal lamella at least twice as large as adjacent palmar scales.
Measurements and scale counts based on right side of animals unless otherwise noted. Scale counts and external
observations of morphology were made using a Nikon SMZ1000 stereo dissecting microscope.
Comparisons were made with museum material in the collections of the Institut Royal des Sciences Naturelles
de Belgique, Brussels (IRSNB), Muséum National d’Histoire Naturelle, Paris (MNHN), Chulalongkorn University
Museum of Zoology (CUMZ), Thailand Natural History Museum, Pathum Thani, Thailand (THNHM]), Royal
Forest Department, Bangkok, Thailand (RFD), Zoological Survey of India, Kolkata (ZSI), and California Academy
of Sciences, San Francisco (CAS) (see Ulber 1993; Pauwels et al. 2000b; Pauwels & Chan-ard 2006; Kunya et al.
2014, 2015; Panitvong et al. 2014; Sumontha et al. 2014, 2015 for specimens examined and sources of data).
Original published descriptions and descriptions provided in broader faunal and taxonomic treatments (e.g., Rösler
& Glaw 2008; Ziegler et al. 2010; Ngo & Grismer 2012; Pauwels & Sumontha 2014; Nguyen et al. 2014; Nazarov
et al. 2014) were also consulted.
Cyrtodactylus phetchaburiensis sp. nov.
Figs. 1–3
Holotype. IRSNB 2682 (formerly IRSNB 16682), adult male (Fig. 1); Thailand, Phetchaburi Province, Tha Yang
District, Klatluang Subdistrict, Khao (= Mountain) Tomo, Tham (= Cave) Khao Tomo (12º47.98’N, 99º44.36’E),
collected by local villagers, September 2003.
Paratype. IRSNB 2683 (formerly IRSNB 16650), adult female (Fig. 2); Thailand, Phetchaburi Province, Tha
Yang District, collector unknown, no date.
Additional specimens. Other Phetchaburi material initially referred to C. oldhami is here tentatively assigned
to Cyrtodactylus phetchaburiensis sp. nov., but with some reservation (see Discussion): THNHM 1247–1248,
Thailand, Phetchaburi Province, Tha Yang District, Kaeng Krachan National Park, along Pala-U Waterfall;
THNHM 1256–1257, 1283–1284, Thailand, Phetchaburi Province, Tha Yang District, Kaeng Krachan National
Park, Khao Phanoen Thung; THNHM 20245–20249 (formerly TNRC 52-3868–3872) (Fig. 3), Thailand,
Phetchaburi Province, Tha Yang District, Kaeng Krachan National Park, near km 23 on the road from Thung Peak
(12°47’N, 99°23’E, 900 m elevation), collected by J. Nabhitabhata, 22 June 1990.
Diagnosis. A medium-sized Cyrtodactylus (SVL to at least 63.2 mm); body slender; limbs and digits relatively
long, slender; original tail longer than SVL; enlarged pair of first postmental scales in contact with one another
behind mental, a smaller pair of enlarged chin shields (second postmentals) lateral to these; small, mostly keeled
tubercles in 20 regular longitudinal rows on dorsum; 33 scales across mid-venter between lowest rows of flank
tubercles; ventrolateral folds weakly developed and atuberculate; enlarged row of femoral scales present; five
precloacal pores in male, femoral pores and precloacal groove absent; 5–6 broad basal lamellae and 11 narrow
distal lamellae beneath digit IV of pes; most of post-pygal portion of tail atuberculate, a single median row of
transversely enlarged subcaudal scales present. Colour pattern of large, dark markings with diffuse edges on a fawn
background extending from shoulders to sacrum. A pair of dark scapular patches and a broad occipital band
outlined by thin white borders. Flanks pale brown with scattered lighter blotches. Tail distinctly banded with
thicker brown annuli, becoming darker posteriorly, alternating with narrower white annuli or bands.
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FIGURE 1. Adult male holotype of Cyrtodactylus phetchaburiensis sp. nov. (IRSNB 2682). Scale bar = 10 mm.
FIGURE 2. Adult female paratype of Cyrtodactylus phetchaburiensis sp. nov. (IRSNB 2683). Scale bar = 10 mm.
Description of holotype. Adult male. SVL 57.5 mm; TailL 70.8 mm (original). Head long (HeadL/SVL ratio
0.31), moderately narrow (HeadW/HeadL = 0.60), not markedly depressed (HeadH/HeadL = 0.40), clearly distinct
from neck. Loreal region weakly inflated, canthus rostralis not prominent. Snout moderate in length (SnEye/HeadL
= 0.38), somewhat acuminate; longer than eye diameter (OrbD/SnEye = 0.65); scales on snout small, oval to
oblong, weakly conical with apex directed posteriorly, somewhat heterogeneous, distinctly larger than those on
crown, interorbital and occipital regions. Eye moderately large (OrbD/HeadL = 0.25); pupil vertical with
crenelated margins; supraciliaries short, minute spines on posterior 60% of scales; anterior supraciliaries largest
and bearing a central keel that becomes less pronounced posteriorly. Ear opening obliquely elliptical, demarcated
posteroventrally by a small fold of skin, large (EarL/HeadL = 0.11); eye to ear distance slightly greater than
diameter of eyes (EyeEar/OrbD = 1.10). Rostral much wider (3.0 mm) than deep (1.9 mm), rostral crease about one
half height of rostral. Two enlarged, rhomboidal supranasals separated by a single, small, pentagonal internasal.
Rostral in contact with first supralabials, nostrils, internasal and supranasals. Nostrils oval, slightly
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posterolaterally-directed, each surrounded by supranasal, rostral, first supralabial, two distinct postnasals, and
posteroventrally by a crescentic nasal, occluded posteromedially by a narial valve. Two to three rows of small
scales separate orbit from supralabials. Mental triangular with acute posteriorly-oriented apex separating anterior
40% of postmentals, wider (2.4 mm) than deep (1.8 mm). A pair of enlarged postmentals in contact behind mental,
each bordered anteromedially by mental, anterolaterally by first infralabial, posterolaterally by an enlarged lateral
chinshield (= second postmental) approximately 40% size of first postmental, and posteriorly by three slightly
enlarged granules. Supralabials to midorbital position 8 (both sides); enlarged supralabials to angle of jaws 12
(right) to 13 (left). Infralabials 11 (both sides). Interorbital scale rows across narrowest point of frontal bone 15; 47
scales between left and right supraciliary rows.
Body slender, elongate (TrunkL/SVL = 0.41) with faintly demarcated ventrolateral folds lacking serrations,
tubercles, or spines. Dorsal scales exclusive of tubercles largely homogeneous, conical with slightly posteriorly-
oriented apices; regularly distributed tubercles (5–8 times size of adjacent scales), most circular, some partly
flattened posteriorly, extending from crown and dorsal postorbital region to tail base, on trunk smallest on flanks,
largest in paravertebral position and over sacrum, smaller tubercles on postorbital region, crown, occiput, and nape;
tubercles conical and erect anteriorly, becoming slightly recumbent and strongly keeled posterior to shoulders, in
20 rows at midbody, typically separated from one another by 1–3 dorsal granules. Ventral scales much larger than
dorsals, smooth, oval subimbricate to imbricate, largest on posterior abdomen and especially in precloacal region
where they are approximately 3–4 times the size of those at midbody and assume a subtriangular shape. Midbody
scale rows across belly to ventrolateral folds ~33. Gular region with homogeneous, smooth, juxtaposed granular
scales.
Three (left) and two (right) precloacal pores separated by a single poreless scale. No femoral pores. No
precloacal groove. Anteroventral scales of thigh much larger than posteroventral and posterior, with a distinct
single row of enlarged femoral scales. Two slightly enlarged, smooth postcloacal spurs, anterior larger than
posterior. Hemipeneal bulges evident. Scales on palm and sole smooth, rounded to oval, flattened to slightly
domed. Scalation on dorsal surfaces of hind limbs and distal fore limb similar to body dorsum with enlarged,
conical, partly keeled tubercles interspersed among smaller scales, tubercles separated from one another by 1–2
small scales; proximal portion of forearm lacking tubercles. Fore and hindlimbs relatively long, slender (ForeaL/
SVL = 0.15; CrusL/SVL = 0.19). Digits relatively short, slender, inflected at interphalangeal joints, especially
antepenultimate joint of longer digits, all bearing robust, slightly recurved claws. Basal subdigital lamellae broad,
squarish to rectangular with rounded corners, distalmost generally largest, without scansorial surfaces (3-4-4-4-4)
both manus, 3-3-3-6-4 left pes, 3-4-4-5-4 right pes); narrow lamellae distal to digital inflection and not including
ventral claw sheath: 7-8-10-10-8 (left manus), 7-8-8-9-8 (right manus), 8-9-11-11-11 (left pes), 7-9-11-11-11 (right
pes); basalmost usually fragmented; no interdigital webbing. Relative length of digits: IV > III > II ~ V > I
(manus); IV ~ V ~ III > II > I (pes).
Original tail, long, subcylindrical, gently tapering to pointed tip; longer than SVL (TailL/SVL = 1.23).
Scales of tail dorsum regular, flat, juxtaposed to weakly imbricate; squarish basally, becoming more
rectangular posteriorly; those on basal segments weakly keeled and with slightly pointed free margins, arranged in
visible but indistinct segments of 7 scale rows basally. Basalmost postpygal segments with a transverse row 6
enlarged, flattened, keeled tubercles, each 3–4 times size of adjacent scales, decreasing to 4 tubercles in next
segment and to 2 in the fifth and sixth postpygal segments, becoming smaller and more weakly keeled posteriorly
and lost in the seventh and more posterior segments. Subcaudal scales much enlarged, medial row forming a single
series of enlarged plates, two per tail segment.
Measurements (in mm; holotype followed by paratype): SVL 57.5/63.2, ForeaL 8.8/9.2, CrusL 11.1/11.9,
TailL 70.8/72.6, TailW 5.4/5.4, TrunkL 23.8/27.8, HeadL 17.8/17.6, HeadW 10.7/11.3, HeadH 7.1/7.5, OrbD 4.4/
4.2, EyeEar 4.8/4.9, SnEye 6.7/6.9, NarEye 4.6/5.2, Interorb 2.4/2.3, EarL 1.9/1.7, Internar 2.1/2.1.
Coloration in ethanol: Dorsum pale fawn with a series of large, darker, diffusely-edged markings from
shoulders to sacrum. A pair of dark scapular patches outlined by a thin white border. Flanks pale brown with
scattered lighter blotches.
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FIGURE 3. Specimens tentatively referable to Cyrtodactylus phetchaburiensis sp. nov. but showing variation in dorsal colour
pattern (see Discussion).
A well demarcated medium brown band with diffuse white margins extending between postorbital regions of
left and right sides across the occiput, band posteriorly bifid across mid-occiput. Crown and snout mid-brown with
some lighter and darker irregular patches. Area below occipital band pale cream, posterior supralabials whitish,
becoming cream with diffuse darker centres anteriorly.
Limbs pale brown, without distinct markings, feet darker than limbs. Venter beige with all scales bearing tiny
brown to black flecks, enlarged precloacal scales paler than remainder of venter. Tail distinctly banded with thicker
brown annuli, becoming darker posteriorly, alternating with narrower white annuli; 13 white bands in total,
basalmost merely dorsal spots, becoming more entire posteriorly, but not extending onto enlarged median
subcaudals, which are brown with a darker free margin.
Variation . The female paratype agrees in nearly all respects of pholidosis with the holotype, but lacks any
trace of precloacal pores and has smaller, less prominent postcloacal spurs. The paratype is, however, more boldly
patterned. The white stripes on the neck and shoulders fully enclose a dark brown patch that is itself bisected by a
white band perpendicular to the long axis of the body. The dark brown blotches on the trunk are discrete and have
irregular margins and two parallel rows along the dorsum, with some sequential blotches partly confluent with one
another. A more diffuse series of dark blotches is present on each flank. The pale background colour forms an
irregular vertebral stripe separating the two paravertebral rows of dark blotches. The crown is distinctly lighter than
the occipital band. All labial scales are whitish, without dark centers. The original tail bears 11 white bands, all
with dark brown margins; those at midportion of tail enclosing dark brown spots or dashes.
Additional material tentatively referred to Cyrtodactylus phetchaburiensis sp. nov. is similar in scalation
features to the holotype, but shows significant variation in dorsal pattern (see Discussion).
Etymology. The specific epithet refers to Phetchaburi Province, south-western Thailand, to which the new
species appears to be endemic.
Distribution and natural history. The types are from the Tha Yang District of Phetchaburi Province (Fig. 4).
The holotype was found by day at about two meters above the ground within a cave-like crevice formed by large
karst boulders in a secondary forest area. Ecological information from other referred specimens, from Kaeng
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Krachan and Tha Yang districts, suggests that the species occurs in moist evergreen forest and climbs on
vegetation-covered walls and tree trunks (Nabhitabhata in Ulber 1993; Pauwels and Chan-ard 2006). Ulber (1993:
195) quoted Jarujin Nabhitabhata’s field notes: ‘‘All specimens collected in moist evergreen forest on the hill,
found at night crawling on vertical steep wayside walls of the forest tracks which are covered with dense vegetation
and roots in cracks on the walls. Their reddish brown colour blends well with the colour of the lateritic soil. A
young was dug out of the rock pile on the forest track’’. It is highly probable that the species will be found on
adjacent hills and mountains covered with primary and mature secondary forests in the northern part of Prachuap
Khiri Khan Province and in the southern part of Ratchaburi Province.
FIGURE 4. Map of the Phetchaburi Province (right) showing the type locality of Cyrtodactylus phetchaburiensis sp. nov.
(white circle with black dot). Other specimens referred to the new species have been collected at localities throughout Kaeng
Krachan National Park, which occupies most of the western half of the province and abuts the border with Myanmar. Inset map
at left shows the relative position of Phetchaburi Province, marked in black.
Comparisons with other species. Wood et al. (2012) established that there was a geographically coherent
pattern of phylogenetic relationships among Cyrtodactylus. The Thai material they sampled did not constitute a
monophyletic group, but fell out in several clusters in a more inclusive clade that included most members of the
genus exclusive of those occurring west of the Salween River. Some groups within this large clade are especially
well-circumscribed geographically and are also generally morphologically distinctive from the new species and
other Thai Cyrtodactylus. These include the members of the Philippine/Bornean clade, the Australo-Papuan clade
and the Lesser Sunda clade (see Wood et al. 2012), as well as the distinctive peninsular Indian/Sri Lankan members
of the subgenus Geckoella Gray (Agarwal & Karanth 2015). However, because Thai, Indochinese, and peninsular
Malaysian taxa are less strictly geographically constrained, it is appropriate to compare the new species with all of
these.
Ngo & Grismer (2012) and Pauwels & Sumontha (2014) provided a summary of Cyrtodactylus from these
regions that have precloacal pores but lack femoral pores, the condition shown by C. phetchaburiensis sp. nov.
Among these taxa, the new species may be distinguished by its possession of enlarged femoral scales from C.
aurensis Grismer, C. bobrovi Nguyen, Le, Pham, Ngo, Hoang, Pham & Ziegler, C. buchardi David, Teynié &
Ohler, C. chauquangensis Hoang, Orlov, Ananjeva, Johns, Hoang & Dau, C. cryptus Heidrich,sler, Vu, Böhme
& Ziegler, C. durio Grismer, Shahrul, Quah, Muin, Chan, Grismer & Norhayati, C. elok Dring, C. hontreensis Ngo,
Grismer & Grismer, C. nigriocularis Nguyen, Orlov & Darevsky, C. otai Nguyen, Le, Pham, Ngo, Hoang, Pham &
Ziegler, C. pageli Schneider, Nguyen, Schmitz, Kingsada, Auer & Ziegler, C. pantiensis Grismer, Chan, Grismer,
Wood & Belabut, C. pseudoquadrivirgatus Rösler, Vu, Nguyen, Ngo & Ziegler, C. spelaeus Nazarov, Poyarkov,
Orlov, Nguyen, Milto, Martynov, Konstantinov & Chulisov, C. stresemanni Rösler & Glaw, C. sumonthai Bauer,
Pauwels & Chanhome, and C. sworderi (Smith).
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By its possession of enlarged subcaudal scales, Cyrtodactylus phetchaburiensis sp. nov. is distinguished from
C. bidoupimontis Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. buchardi, C. bugiamapensis
Nazarov, Poyarkov, Orlov, Phung, Nguyen, Hoang & Ziegler, C. cattienensis Geissler, Nazarov, Orlov, Böhme,
Phung, Nguyen & Ziegler, C. chauquangensis, C. cryptus, C. cucdongensis Schneider, Phung, Le, Nguyen &
Ziegler, C. durio, C. irregularis (Smith), C. martini Ngo, C. metropolis Grismer, Wood, Onn, Anuar & Muin, C.
pantiensis, C. papilionoides Ulber & Grossmann, C. payacola Johnson, Quah, Anuar, Muin, Wood, Grismer,
Greer, Chan, Norhayati, Bauer & Grismer, C. pseudoquadrivirgatus, C. quadrivirgatus Taylor, C. stresemanni, C.
sworderi, C. vilaphongi Schneider, Nguyen, Duc Le, Nophaseud, Bonkowski & Ziegler, and C. ziegleri Nazarov,
Orlov, Nguyen & Ho.
By its possession of 20 longitudinal rows of dorsal tubercles at midbody, Cyrtodactylus phetchaburiensis sp.
nov. is distinguishable from C. buchardi (25), C. eisenmanae Ngo (14), C. elok (5–10), C. hontreensis (14), C.
nigriocularis (0), C. pageli (9–14), C. papilionoides (12–14), C. spelaeus (10), C. stresemanni (13), C. sumonthai
(12), and C. wangkulangkulae Sumontha, Pauwels, Suwannakarn, Nutatheera & Sodob (10).
Among remaining species C. phetchaburiensis sp. nov. is distinguished by a lower number of lamellae beneath
digit IV of the pes (16–17) from C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi
& Dangsri (19), C. leegrismeri
Chan & Norhayati (18–20), C. puhuensis Nguyen, Yang, Le, Nguyen, Orlov,
Hoang, Nguyen, Jin, Rao, Hoang Che, Murphy & Zhang (23), C. samroiyot Pauwels & Sumontha (20), C. sanook
Pauwels, Sumontha, Latinne & Grismer (20), C. surin Chan-ard & Makchai (18), C. teyniei David, Nguyen,
Schneider & Ziegler (19–20), C. wayakonei Nguyen, Kingsada, Rösler, Auer & Ziegler (19–20); and by a lower
number of precloacal pores (5) than C. angularis (Smith) (6), C. doisuthep (6), C. intermedius (Smith) (8–10), C.
phuquocensis Ngo, Grismer & Grismer (7–9), C. samroiyot (7), C. teyniei (14 in female holotype) and C.
wayakonei (6–8), and a greater number of precloacal pores than C. paradoxus Darevsky & Szczerbak (0–4), and C.
thuongae Phung, Van Schingen, Ziegler & Nguyen (0–1).
In addition, the colour pattern of C. phetchaburiensis sp. nov. is distinctly different from all Thai, Malay, and
Indochinese Cyrtodactylus that share the lack of femoral pores, presence of six or fewer precloacal pores, enlarged
femoral scales present, broad subcaudal scutes, and 19–24 dorsal rows of tubercles (or condition unknown).
Specifically, it differs from C. condorensis (Smith), C. doisuthep, C. leegrismeri, C. puhuensis, C. saiyok
Panitvong, Sumontha, Tunprasert & Pauwels, C. samroiyot, C. sanook, C. thochuensis Ngo & Grismer, C.
thuongae and C. yangbayensis Ngo & Chan in lacking body bands or well-defined transverse markings, from C.
angularis in lacking W-shaped dorsal markings, from C. wayakonei in lacking a reticulated pattern on the head and
in possessing a well-defined nuchal loop, from C. paradoxus in lacking a thin, pale vertebral stripe and in having
an entire (versus bisected) nuchal loop, from C. peguensis in lacking a reticulated pattern on the head and discrete
dark dorsal spots separated by narrow pale margins or reticulations, and from C. oldhami in having a diffuse pattern
of dark blotches on a light background rather than four longitudinal series of white spots. In addition, C.
phetchaburiensis sp. nov. differs from all of these species is having discrete, dark scapular patches with thin white
borders.
Discussion
Cyrtodactylus phetchaburiensis sp. nov. is most similar to C. oldhami and C. peguensis, which, among sequenced
members of the genus, are each others closest relatives (Wood et al. 2012). These in turn are sister to the
morphologically divergent C. tigroides Bauer, Sumontha & Pauwels from nearby Kanchanaburi Province. The
clade as a whole extends along the Thai-Myanmar border and south of the Isthmus of Kra deep into the Thai-Malay
Peninsula. The distribution of both C. oldhami and C. peguensis in Thailand is poorly documented. Chan-ard et al.
(2015) suggested that the former extends from Kanchanaburi south and throughout much of western peninsular
Thailand, whereas the latter species is more restricted to Kanchanaburi and Ratchaburi, with scattered populations
in evergreen forests of peninsular Thailand (Taylor 1963; Pauwels et al. 2000b). The Myanmar distribution of these
two taxa is even more poorly known (Bauer 2002).
Phetchaburi specimens of Cyrtodactylus oldhami from Kaeng Krachan National Park, Tha Yang District, have
previously been discussed by Ulber (1993). The scale counts for these specimens approximate those of C.
phetchaburiensis sp. nov., with dorsal tubercle rows 19–24, ventral scales 30–33, and four precloacal pores. The
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colour patterns of these specimens, however, differ substantially from the types of the new species (Fig. 3). The
juvenile specimen (THNHM 20249 = TNRC 52-3872) exhibits a pattern very typical of true C. oldhami, with lines
of small, separate white spots. A gravid female (THNHM 20247 = TNRC 52-3869) has more-or-less continuous
lines anteriorly, breaking into separated spots or dashes posteriorly. The remaining specimens, all adult males
(THNHM 20245 = TNRC 52-3868, THNHM 20248 = TNRC 52-3870, THNHM 20246 = TNRC 52-3871), have
more-or-less continuous pairs of dorsal stripes, the paravertebral pair enclosing an area of lighter colour than the
remainder of the dorsum.
FIGURE 5. Holotype of Cyrtodactylus oldhami (Theobald), ZSI 5858, showing the pattern of four longitudinal series of white
spots characteristic of this species.
The relationship between the pattern elements in the types of C. phetchaburiensis sp. nov. and the Kaeng
Krachan specimens is clear. The scapular patches of the new species represent a much truncated version of the
anterior stripes of the other specimens. It seems unlikely that two members of the C. oldhami group that are so
similar would occur in sympatry or parapatry in the Tha Yang District, suggesting that the specimens reported upon
by Ulber (1993), and possibly others (Pauwels & Chan-ard 2006), are conspecific. Nonetheless, the colour pattern
differences are significant and until further data, including molecular data, are available, our assignment of the
fully striped specimens to C. phetchaburiensis sp. nov. remains tentative. It should be noted that the caption of the
Figure 7 in Ulber (1993: 194) erroneously mentions ‘Cyrtodactylus interdigitalis sp. nov.’’, whereas his Figure 4
(loc. cit.: 189, showing the type series of C. interdigitalis Ulber, 1993) says ‘Cyrtodactylus oldhami (Theobald)’’.
Ulbers Figure 8 (loc. cit.: 194), showing the cloacal area and the four precloacal pores of the male TNRC 52-3870
(= field nr JN 8075) is correctly labeled.
The taxonomic revision of the C. oldhami complex is hindered by the fact that the provenance of the type is
unknown (Smith 1935; Taylor 1963; Das et al. 1998). However, its colour pattern remains visible (Annandale
1913; Taylor 1963; Fig. 5) confirming that it is consistent with the application of the name to the familiar form with
a series of four lines of spots on the dorsum, which occurs both WNW of Phetchaburi in the Dewei District of the
Tanintharyi Division of Myanmar (CAS 229790, 245740, 247959, 247979, 24799:) and south of it, in the
Kawthaung District of the Tanintharyi Division of Myanmar (CAS 247475) as well as in Thai peninsular localities
(see, for example, illustrations of specimens from Chumphon in Manthey & Grossmann 1997, Chan-ard et al.
1999).
Phetchaburi is one of the most well studied Thai provinces (Pauwels et al. 2000a, 2003, 2009; Pauwels &
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Kheowyoo 2004; Pauwels & Chan-ard 2006). Pauwels et al. (2009) reported that 102 reptile species were known
from Phetchaburi Province. Since then, additional species recorded have included a new species of Ptychozoon
(Sumontha et al. 2012). Although C. phetchaburiensis sp. nov. adds another putative endemic to the provincial
total, true C. oldhami likely needs to be removed from the Phetchaburi species list, which, among Cyrtodactylus,
includes only C. brevipalmatus (Ulber 1993; Manthey & Grossmann 1997; Pauwels et al. 2003) in addition.
Acknowledgements
We thank Kumthorn Thirakhupt (CUMZ, Bangkok), Georges Lenglet and Sébastien Bruaux (IRSNB) for allowing
us to study the type and reference material. Michael Cota (THNHM) kindly provided photographs of the series of
specimens from Kaeng Krachan and Indraneil Das provided the photo of the holotype of Cyrtodactylus oldhami.
AMB was supported by grants DEB 0844523 and EF-1241885 (subaward 13-0632) from the National Science
Foundation of the United States.
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... The Cyrtodactylus oldhami group is restricted to a narrow geographic range on the Thai-Malay Peninsula and Myanmar northward into Kanchanaburi Province in western Thailand (Panitvong et al. 2014;Pauwels et al. 2016;Connette et al. 2017;Grismer et al. 2018bGrismer et al. , 2021b. The oldhami group is one of the most taxonomically diverse species groups of Cyrtodactylus in Thailand. ...
... The group is monophyletic and contains at least seven nominal species (Grismer et al. 2021b), of which five occur in Thailand, i.e., C. oldhami (Theobald, 1876), C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels, 2014, C. sanook Pauwels, Sumontha, Latinne & Grismer, 2013, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome, 2004, and C. zebraicus Taylor, 1962. Cyrtodactylus phetchaburiensis Pauwels, Sumontha & Bauer, 2016 from southern Thailand may also belong to the oldhami group, based on morphological characters (Pauwels et al. 2016), but this hypothesis remains untested by molecular data. Two additional species in the group are known from Myanmar, i.e., C. lenya Mulcahy, Thura &Zug, 2017, andC. ...
... The group is monophyletic and contains at least seven nominal species (Grismer et al. 2021b), of which five occur in Thailand, i.e., C. oldhami (Theobald, 1876), C. saiyok Panitvong, Sumontha, Tunprasert & Pauwels, 2014, C. sanook Pauwels, Sumontha, Latinne & Grismer, 2013, C. thirakhupti Pauwels, Bauer, Sumontha & Chanhome, 2004, and C. zebraicus Taylor, 1962. Cyrtodactylus phetchaburiensis Pauwels, Sumontha & Bauer, 2016 from southern Thailand may also belong to the oldhami group, based on morphological characters (Pauwels et al. 2016), but this hypothesis remains untested by molecular data. Two additional species in the group are known from Myanmar, i.e., C. lenya Mulcahy, Thura &Zug, 2017, andC. ...
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... New species are continuously being described each year through fieldwork in underexplored localities and the development of molecular techniques, especially DNA barcoding (Hebert et al., 2003). A significant number of Cyrtodactylus have been described from tropical karst forests in regions of Southeast Asia, including Myanmar, Malaysia, Laos, Vietnam, Thailand, Cambodia, Indonesia, Singapore, Timor Leste, Philippines, Brunei and China (e.g., Pauwels et al., 2016;Sitthivong et al., 2019;Grismer et al., 2020). Four species of Cyrtodactylus are currently recognized in China (Wang et al., 2020) including C. cayuensis (Li, 2007;Agarwal et al., 2018), C. tibetanus (Boulenger, 1905), C. zhaoermii (Shi and Zhao, 2010) and C. wayakonei (Nguyen et al., 2010). ...
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Cyrtodactylus geckos are one of the most speciose and diverse groups of extant lizards known, distributed throughout the Asian and Pacific realms. Using molecular phylogenetic methods and supporting morphological data, we describe a new species of Cyrtodactylus in Daweishan National Nature Reserve, Yunnan Province, China. Cyrtodactylus hekouensis sp. nov. can be morphologically distinguished from its nearby congeners by the following characters: maximum SVL 92.3 mm and TL 98.5 mm; 11-12 supralabials; 11-12 infralabials; 36-57 scale rows between the fifth supralabials; 10-13 dorsal tubercles rows; 3 postnasals on blunt and smooth front snout; precloacal-femoral pores in a continuous series of 33-39 (females with pitted scales) located under vent/cloaca and thighs in both sexes; precloacal groove absent; 3/3 postcloacal tubercles; subdigital lamellae under the fourth finger 21 or 22, under the fourth toe 20-23; smooth midbody with smooth venter and tuberculate dorsal scale rows, tubercles from head to tail base; dorsal transverse patterns are generally large, bilaterally symmetrical. The results of the phylogenetic analysis recover specimens of this new species as sister to a clade containing C. wayakonei and C. martini. Uncorrected pairwise intraspecific distances were < 1%, and distances between our new species and other Cyrtodactylus species from nearby countries ranged from 14.2% to 26.8%.
... nov. with other congeners from Vietnam and neighboring countries in the mainland Indochina region, including Laos, Cambodia, and Thailand based on examination of specimens (see Appendix) and data from literature (Luu et al. 2014;Nazarov et al. 2014;Nguyen et al. 2014;Panitvong et al. 2014;Schneider et al. 2014 a,b;Nurngsomsri et al. 2014;Grismer et al. 2015;Luu et al. 2015;Sumontha et al. 2015;Pauwels et al. 2016;Le et al. 2016;Luu et al. 2016aLuu et al. ,b,c, 2017Nguyen et al. 2017;Chuaynkern et al. 2018;Pauwels et al. 2018;Nazarov et al. 2018;Murdoch et al. 2019;Pham et al. 2019;Sitthivong et al. 2019;Schneider et al. 2020;Ostrowski et al. 2020Ostrowski et al. , 2021. The new species can be differentiated from other known species of the genus Cyrtodactylus by morphological characters (see Table 3). ...
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A new species of the genus Cyrtodactylus is described from Dien Bien Province, northwestern Vietnam based on morphological and molecular data. Cyrtodactylus ngati sp. nov. can be distinguished from remaining congeners by the following combination of characters: maximum SVL 69.3 mm; dorsal pattern consisting of six dark irregular transverse bands between limb insertions; inter-supranasals one; dorsal tubercles present on occiput, body, hind limbs and on first half of tail; 17-22 irregular dorsal tubercle rows at midbody; lateral folds clearly defined, with interspersed tubercles; 32-38 ventral scales between ventrolateral folds; 13 precloacal pores separated by a diastema of 5/5 poreless scales from a series of 7/7 femoral pores in enlarged femoral scales; precloacal and femoral pores absent in females; 1-3 postcloacal tubercles on each side; transversely enlarged median subcaudal scales absent. In the molecular analyses, the new species is shown to be the sister taxon to C. interdigitalis from Thailand. This is the 47 th species of the genus Cyrtodactylus and the first member of the C. brevipalmatus species group recorded from Vietnam.
... The present study reveals an underestimated diversity of Cytrodactylus geckos within the Andaman and Nicobar archipelago until now, which is in line with the vast number of new species discoveries made throughout most parts of the range of this genus (e.g. Pauwels et al. 2016;Oliver et al., 2019;Quah et al., 2019;Grismer et al., 2015;Zeigler et al., 2013;Grismer et al., 2008;Grismer et al., 2018b, Nazarov1 et al., 2018Grismer et al., 2019). Since Blyth (1861) described C. rubidus from the Andaman Islands, there had been a vast gap of 136 years in-between until the description of the second species, C. adleri by Das (1997). ...
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A comprehensive review of members of the gekkonid genus Cyrtodactylus Gray, 1827 from the Andaman and Nicobar Islands was undertaken to assess the true diversity of the genus in this region. Samples collected across different islands within the archipelago show significant and consistent morphological variation associated with the region of origin. Detailed redescriptions are presented for the two known species Cyrtodactylus rubidus and C. adleri. Two new species C. nicobaricus sp. nov. and C. camortensis sp. nov. are described from the northern and central group of islands of the Nicobar archipelago based on morphological distinction and geographic separation from the above two named species from this archipelago. Information on geographic distribution, natural history and conservation status for all of these species are presented.
... nov. with other congeners from Laos and neighboring countries in the mainland Indochina region, including Vietnam, Cambodia, and Thailand based on examination of specimens (see Appendix) and data from literature (Luu et al. 2014;Nazarov et al. 2014;Nguyen et al. 2014;Panitvong et al. 2014;Schneider et al. 2014 a,b;Luu et al. 2015;Nguyen et al. 2015b;Sumontha et al. 2014Sumontha et al. , 2015Pauwels et al. 2016;Le et al. 2016;Luu et al. 2016 a,b,c;Luu et al. 2017;Nguyen et al. 2017;Pauwels et al. 2018;Nazarov et al. 2018;Murdoch et al. 2019;Pham et al. 2019). The new species can be differentiated from other known species of the genus Cyrtodactylus by morphological characters (see Table 3). ...
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A new species of the genus Cyrtodactylus from Vientiane Province, northern Laos is described based on morphological and molecular data. Cyrtodactylus muangfuangensis sp. nov. can be distinguished from remaining congeners by the following combination of characters: maximum SVL 83.9 mm; dorsal pattern consisting of dark nuchal loop, nape band and five dark transversal bands between limb insertions; intersupranasals two; dorsal tubercles present on occiput, body, hind limbs and tail base; 15 or 16 irregular dorsal tubercle rows at midbody; lateral folds clearly defined, without interspersed tubercles; 31-37 ventral scales between ventrolateral folds; six precloacal pores and 15 femoral pores in males, which are interrupted by six to eight poreless scales; six precloacal pitted scales plus in total 10-15 pitted femoral scales in females, which are separated by six to eight poreless scales; enlarged precloacal and femoral scales present; two or three postcloacal tubercles; median subcaudal scales transversely enlarged. In molecular analyses, the new species is strongly supported as a member of the Cyrtodactylus phongnhakebangensis species group, and weakly corroborated as a sister taxon to C. pageli. Pairwise genetic comparison shows that it is at least 18% divergent from other congeners in the species group based on a fragment of the mitochondrial cytochrome C oxidase subunit I gene.
... The number of described species continues to rise with many recent Cyrtodactylus descriptions of species inhabiting limestone-karst ecosystems. Between 2016 and 2018, 34 new Cyrtodactylus species were described, of which 27 are endemic to karst ecosystems (Agarwal 2016;Grismer et al. 2016a;2018a;Luu et al. 2016a;Mecke et al. 2016;Nazarov et al. 2018;Nguyen et al. 2017;Nielsen & Oliver 2017;Oliver et al. 2016;Pauwels et al. 2016;Riyanto et al. 2017). Despite the high number of Cyrtodactylus species recorded and the rapid rate at which new species are being described, the island of Borneo is largely underrepresented with a mere eight documented species (Das 2005). ...
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The island of Borneo lies within one of the most biodiverse regions in the world. Despite this, its documented gekkonid diversity is not commensurate with other areas of Southeast Asia. The megadiverse genus Cyrtodactylus is especially un-derrepresented. Limestone-karst ecosystems, in particular, harbor many endemic Cyrtodactylus species, but only one karst-dwelling species is currently recognized from Borneo. This paper adds two additional karst-dwelling Cyrtodactylus species-C. muluensis sp. nov. and C. limajalur sp. nov.-from Sarawak, Malaysia. Cyrtodactylus muluensis sp. nov. is endemic to Gunung Mulu and is distinguished from its congeners by having a precloacal groove, 31-38 ventral scales, a maximum SVL of at least 88 mm, enlarged subcaudals, 19-20 subdigital lamellae, and a banded dorsal body pattern. Cyr-todactylus limajalur sp. nov. is endemic to the Serian region and is distinguished from its congeners by having 33-42 ven-tral scales, enlarged subcaudals, a precloacal pit, a maximum SVL of at least 94 mm, 5-6 enlarged femoral scales, 19-22 subdigital lamellae, and five distinct bands on the dorsum. Both species are phylogenetically distinct and deeply divergent from all other congeners. The description of two new karst-dwelling species highlights the need to conserve karst habitats and the endemic species they harbor.
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We present the first multilocus molecular phylogeny focused on mangrove pit vipers of the Trimeresurus erythrurus‐purpureomaculatus complex based on novel topotypic material and expanded geographic sampling. Previously inferred paraphyly of T. purpureomaculatus was resolved and our results demonstrate distinct phylogeographic patterns that are latitudinally stratified and consistent with isolation‐by‐distance and isolation‐by‐environment. A clear genetic break is detected at the Isthmus of Kra biogeographic divide and niche overlap among major genetic clades is limited. The association between colour polymorphisms and phylogeographic structure suggests that contemporary or historical introgression between T. erythrurus and T. purpureomaculatus could have occurred. This study provides a clear roadmap to guide future genomic research to improve our understanding of this charismatic, yet poorly studied group of snakes.
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A new species of Cyrtodactylus from Tak Province, Thailand, Cyrtodactylus amphipetraeus sp. nov., is described using an integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene NADH dehydrogenase subunit 2 (ND2). The phylogenetic analyses place the new species within the C. sinyineensis group which was previously thought to be endemic to the Salween Basin in southern Myanmar. The phylogeny also places C. inthanon in the C. sinyneensis group which is expanded herein to also include the group’s sister species C. doisuthep. Along with C. amphipetraeus sp. nov., these are the first three species of the C. sinyineensis group to be found outside of Myanmar east of the Tenasserim Mountains. The Tenasserim Mountain region is discussed as an area of cladogeneic turnover.
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We describe a new species of Cyrtodactylus on the basis of four specimens collected from Phu My, Binh Dinh Province, southern Vietnam. Cyrtodactylus phumyensis sp. nov. is distinguished from the remaining Indochinese bent-toed geckos by a combination of the following characters: size small (SVL up to 66.8 mm); two internasals; dorsal tubercle rows 18 or 19 at midbody; ventral scale rows 33-41; ventrolateral folds slightly developed; each thigh with 5-7 enlarged femoral scales; femoral pores absent in males and female; a series of 5-7 precloacal pores plus a pitted, enlarged precloacal scale in males; 6 pitted, enlarged precloacal scales in female; paravertebral tubercles 20-23; lamellae under toe IV 18-21; small subcaudal scales, not transversely enlarged; two irregular dark longitudinal stripes on shoulder. In phylogenetic analyses, the new species is recovered as a member of the Cyrtodactylus irregularis species group, and strongly supported as a sister taxon of C. cucdongensis from Khanh Hoa Province.
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An integrative taxonomic analysis of the distantly related Cyrtodactylus condorensis and intermedius species complexes of the Mekong Delta revealed that C. paradoxus is a junior synonym of C. condorensis and that C. thochuensis is a junior synonym of C. leegrismeri. Additionally, the analysis revealed that a cave-dwelling ecomorpholgy has evolved independently early on in the evolution of both complexes (represented by C. hontreensis in the intermedius complex and C. grismeri and C. eisenmani in the condorensis complex) and cave ecomorphs exist in sympatry-but not syntopy-with general scansorial ecomorphs. Multiple, recent, cyclical, glacioeustatic driven changes in sea levels across the Sunda Shelf are hypothesized to account for the evolution and distribution of the widely separated, conspecific insular populations of C. condorensis and C. leegrismeri. The independent evolution of cave ecomorphology is proposed to have been driven by competition avoidance. Habitat islands across the Mekong Delta are an important source of endemism and in need of protection.
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A new cave-dwelling species of Cyrtodactylus Gray is described from Mae Hong Son Province in northern Thailand. The new species, C. erythrops sp. nov., is characterized by its moderate size (snout-vent length to at least 78 mm), relatively large, closely-spaced, flattened tubercles in 18–20 irregular rows at midbody, low number of ventral scales across midbody (28), absence of precloacal groove, presence of precloacal and femoral pores separated by a diastema, broad subcaudal plates, and dorsal pattern of dark spots and blotches. It is the fifth species of cave-dwelling Cyrtodactylus recorded from Thailand and its discovery adds to the mounting evidence that this genus exhibits unprecedented levels of localized endemism throughout tropical Southeast Asia.
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We describe a new gekkonid lizard from Batang Padang, Perak province, Malaysia, based on a single specimen collected almost ICK) years ago. Cyrtodactylus stresemanni sp. nov. apparently differs from all other species in the genus by large tubercles on the ventral side of the tail, suggesting an isolated position within the genus. A literature survey of meristic and mensural data of all described Cyrtodactylus species revealed further diagnostic characters to distinguish C. stresemanni from all other species.
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Two new cave-dwelling species of Cyrtodactylus from mainland southwestern Vietnam and an offshore island are differentiated from all other congeners in lacking a precloacal groove and precloacal pores, presence of enlarged femoral scales beneath thigh, and in having a color pattern consisting of four or five narrow white bands on the body dorsum and one on the tail. Cyrtodactylus grismeri sp. nov. is reddish brown, has a mean SVL of 87.6 ± 3.8 mm (n = 9), 18-22 irregular longitudinal rows of weakly-keeled tubercles at midbody between the lateral folds, 33-38 ventral scales between ventrolateral folds, 0-3 enlarged scales beneath thighs, and 20-24 subdigital lamellae under the first toe. Cyrtodactylus eisenmani sp. nov. is chocolate brown, has a mean SVL of 81.3 ± 5.0 mm (n = 5), 14 irregular longitudinal rows of weakly keeled tubercles at midbody between ventrolateral folds, 44-45 ventral scales between ventrolateral folds at midbody, 4-6 enlarged femoral scales beneath each thigh, and 22-25 subdigital lamellae under the first toe. This discovery increases the total number of Cyrtodactylus found in Vietnam to fifteen.