AnimaliaSquamataGekkonidaeTermprayoonKorkhwanRujirawanAttapolGrismerL. LeeJr.Perry L. WoodAowpholAnchaleeTaxonomic reassessment and phylogenetic placement of Cyrtodactylus phuketensis (Reptilia, Gekkonidae) based on morphological and molecular evidenceZookeys285202110409112110.3897/zookeys.1040.65750D765E47B-9F30-541B-8C53-5E2EC6C7D3F3 Cyrtodactylus macrotuberculatus Grismer and Ahmad, 2008Figure 4 Cyrtodactylus macrotuberculatusGrismer & Ahmad, 2008: 55; Grismer 2011: 406; Grismer et al. 2012: 45. Cyrtodactylus phuketensisSumontha et al., 2012: 62.Type specimens.

Holotype (adult male, ZRC 2.6754) from Malaysia, Kedah, Pulau Langkawi, Gunung Raya; Paratypes: Malaysia, Kedah, Pulau Langkawi, Gunung Raya: ZRC 2.6755–2.6756, Telaga Tujuh: ZRC 2.6757, Lubuk Semilang: ZRC 2.6758.

Additional specimens examined

(including types of C. phuketensis). Malaysia – Kedah, Pulau Langkawi, Gunung Raya: LSUHC 09428–09429, LSUHC 09432; Perlis, Perlis State Park: LSUHC 09981, LSUHC 10097, ZRC 2.4869. Thailand – Satun Province, Mueang Satun District, Adang Island: ZMKU R 00871–00875, ZMKU R 00879–00882, Rawi Island: ZMKU R 00883–00889; Songkhla Province, Hat Yai District, Chalung Sub-district: ZMKU R 00876–00878; Phuket Province, Thalang District: PSUZC-RT 2010.58, THNHM 15378, ZMKU R 00894–00896, Kathu District: ZMKU R 00890–00893, ZMKU R 00897–00898 (Table 1).

Expanded diagnosis.

Cyrtodactylus macrotuberculatus can be separated from all other species of C. pulchellus complex by having the following combination of characters (Table 6): (1) maximum SVL 117.9 mm (mean 105.0 ± SD 9.8, N = 39); (2) 9–13 supralabial and 7–11 infralabial scales; (3) prominent tuberculation on body; (4) tubercles on ventral surface of forelimbs, gular region, in ventrolateral body folds, and anterior one-third of tail; (5) 34–49 paravertebral tubercles; (6) 19–27 longitudinal tubercle rows; (7) 17–28 ventral scales; (8) 19–23 subdigital lamellae on the fourth toe; (9) 28–42 femoroprecloacal pores in males; (10) deep precloacal groove in males; (11) three or four dark dorsal body bands; (12) body band wider than interspace; (13) 7–10 (N = 12) ringed dark caudal bands on original tail; (14) white caudal bands infused with dark pigmentation in adults; (15) posterior portion of tail in hatchlings and juveniles bands not white.

Morphological measurement (mm), meristic and non-meristic data from males and females of Cyrtodactylus macrotuberculatus. Morphological character abbreviations are defined in the Materials and methods.

CharactersAdult males (N = 24)Adult females (N = 15)All (N = 39)
Mean ± SD(Min–Max)Mean ± SD(Min–Max)Mean ± SD(Min–Max)
SVL 105.7 ± 9.0(88.9–117.9)103.9 ± 11.3(84.1–117.6)105.0 ± 9.8(84.1–117.9)
TW 9.5 ± 1.0(7.9–11.6)8.2 ± 1.5(5.9–10.5)9.0 ± 1.4(5.9–11.6)
FL 17.2 ± 1.6(14.2–19.0)16.8 ± 1.8(13.3–19.3)17.1 ± 1.7(13.3–19.3)
TBL 20.4 ± 1.9(17.3–23.5)20.0 ± 2.1(16.3–19.3)20.2 ± 2.0(16.3–23.5)
AG 50.7 ± 4.9(41.3–58.6)51.4 ± 5.4(42.0–60.2)51.0 ± 5.0(41.3–60.2)
HL 29.4 ± 2.3(24.6–33.3)28.5 ± 3.3(22.8–32.3)29.0 ± 2.7(22.8–33.3)
HW 20.0 ± 2.0(16.3–22.9)18.9 ± 2.0(15.6–21.4)19.6 ± 2.0(15.6–22.9)
HD 12.1 ± 1.4(9.7–14.2)11.3 ± 1.4(9.0–13.5)11.8 ± 1.4(9.0–14.2)
ED 6.8 ± 0.6(5.6–7.9)6.7 ± 0.8(5.2–7.5)6.8 ± 0.7(5.2–7.9)
EE 8.6 ± 1.0(6.5–9.8)8.5 ± 0.9(7.0–9.6)8.6 ± 0.7(5.2–7.9)
ES 11.7 ± 1.0(10.0–13.6)11.5 ± 1.3(9.2–13.1)11.6 ± 1.1(9.2–13.6)
EN 8.7 ± 0.7(7.3–9.8)8.6 ± 1.0(6.8–9.8)8.6 ± 0.8(6.8–9.8)
IO 5.0 ± 0.6(4.0–6.3)4.8 ± 0.8(3.4–5.7)4.8 ± 0.7(3.4–6.3)
EL 2.3 ± 0.3(1.4–2.8)2.3 ± 0.4(1.5–3.0)2.3 ± 0.4(1.4–3.0)
IN2.2 ± 0.5(1.7–2.7)2.1 ± 0.5(1.3–2.9)2.2 ± 0.4(1.3–2.9)
HL/SVL0.28 ± 0.01(0.27–0.30)0.27 ± 0.01(0.26–0.29)0.28 ± 0.01(0.26–0.30)
HW/HL0.68 ± 0.03(0.62–0.74)0.67 ± 0.02(0.62–0.70)0.67 ± 0.03(0.62–0.74)
HD/HL0.41 ± 0.02(0.37–0.45)0.40 ± 0.01(0.38–0.42)0.41 ± 0.02(0.37–0.45)
ES/HL0.40 ± 0.01(0.37–0.41)0.40 ± 0.00(0.39–0.41)0.40 ± 0.01(0.37–0.41)
ED/HL0.23 ± 0.01(0.21–0.27)0.23 ± 0.01(0.22–0.25)0.23 ± 0.01(0.21–0.27)
EL/HL0.08 ± 0.01(0.05–0.10)0.08 ± 0.01(0.05–0.10)0.08 ± 0.01(0.05–0.10)
AG/SVL0.48 ± 0.02(0.43–0.51)0.50 ± 0.01(0.47–0.52)0.49 ± 0.02(0.43–0.52)
FL/SVL0.16 ± 0.00(0.15–0.17)0.16 ± 0.00(0.16–0.17)0.16 ± 0.00(0.15–0.17)
TBL/SVL0.19 ± 0.01(0.18–0.21)0.19 ± 0.00(0.18–0.20)0.19 ± 0.01(0.18–0.21)
TL/SVL1.29 ± 0.04(1.23–1.35)1.27 ± 0.04(1.24–1.34)1.28 ± 0.04(1.23–1.35)
Supralabials9–139–129–13
Infralabials7–117–117–11
TuberculationProminentProminentProminent
Tubercles on ventral surface of forelimbsYesYesYes
Tubercles in gular regionYesYesYes
Ventrolateral fold tuberculateYesYesYes
Paravertebral tubercles37–4934–4734–49
Longitudinal rows of tubercles19–2719–2619–27
Ventral scales17–2819–2617–28
4th toe lamellae19–2319–2319–23
Femoroprecloacal pores28–42No28–42
Precloacal depressionYesNoOnly in males
No. of body bands3 or 43 or 43 or 4
Body band/interspace ratio0.95–1.751.03–1.620.95–1.74
Dark caudal bands on original tail7–97–107–10
Description of adult males.

SVL of adult males range from 88.9–117.9 mm (mean 105.7, N = 24); head moderate in length (HL/SVL 0.27–0.30), width (HW/HL 0.62–0.74), somewhat flattened (HD/HL 0.37–0.45), distinct from neck, triangular in dorsal profile; lores concave; frontal and prefrontal regions deeply concave; canthus rostralis sharply rounded; snout elongate (ES/HL 0.37–0.41), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.21–0.27); ear opening elliptical, moderate in size (EL/HL 0.05–0.10) obliquely oriented; eye to ear distance greater than diameter of eye; rostral rectangular, divided dorsally by an inverted Y or I-shaped furrow, bordered posteriorly by large left and right supranasals and small internasal, bordered laterally by external nares and first supralabials; external nares bordered anteriorly by rostral, dorsally by one large anterior supranasal, posteriorly by two postnasals, ventrally by first supralabial; 9–13 rectangular supralabials extending to just beyond upturn of labial margin, tapering abruptly below midpoint of eye; 7–11 infralabials not tapering in size posteriorly; scales of rostrum and lores slightly raised, larger than granular scales on top of head and occiput, those on posterior portion of canthus rostralis slightly larger; scales on top of head and occiput intermixed with enlarged tubercles; large, boney frontal ridges bordering orbit confluent with boney, transverse, parietal ridge; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by left and right trapezoidal postmentals that contact medially for 40–50% of their length posterior to mental; single row of slightly enlarged, elongate sublabials extending posteriorly to 5th–7th infralabial; small, granular, gular scales intermixed with numerous large, conical tubercles grading posteriorly into larger, conical tubercles on throat which abruptly transition into large, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.43–0.51) with well-defined, tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with large, trihedral, regularly arranged, keeled tubercles separated by no more than three granules at their base; tubercles extend from top of head onto approximately one-half of tail but not onto regenerated tail; tubercles on occiput and nape relatively small, those on body largest; approximately 19–27 longitudinal rows of dorsal tubercles at the mid body; approximately 37–49 paravertebral tubercles; 17–28 flat, imbricate, ventral scales and much larger than dorsal scales; precloacal scales large, smooth; deep precloacal groove (= depression).

Forelimbs moderate in stature, relatively short (FL/SVL 0.15–0.17); virtually no granular scales on dorsal surface of forelimbs, only large, trihedral, keeled tubercles; palmar scales slightly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae nearly square proximal to joint inflection, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.18–0.21), virtually no granular scales on dorsal surfaces of hind limbs, only large, trihedral, keeled tubercles; ventral scales of thigh flat, smooth, imbricate; ventral, tibial scales flat, imbricate, slightly keeled; two rows of enlarged, flat, imbricate, femoroprecloacal scales extend from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 28–42 contiguous, pore-bearing femoroprecloacal scales forming an inverted T bearing a deep, precloacal groove; eight to eleven pores bordering groove; postfemoral scales immediately posterior to the pore-bearing scale row conical, forming an abrupt union on posteroventral margin of thigh; plantar scales low, slightly rounded; digits well-developed, inflected at basal, interphalangeal joints; subdigital lamellae proximal to joint inflection nearly square, only slightly expanded distal to inflection; digits more narrow distal to joints; claws well-developed, sheathed by a dorsal and ventral scale; 19–23 subdigital lamellae on the 4th toe.

Original tail (TL/SVL) moderate in proportions, 123–135% of SVL (mean 128, N = 12), 7.9–11.6 mm in width at base, tapering to a point; dorsal scales at base of tail square, smooth, flat, subimbricate, lacking tubercle on regenerated tail; median row of transversely enlarged, subcaudal scales; shallow caudal furrow; two to five small, postcloacal tubercles at base of tail on hemipenial swellings; all postcloacal scales flat, large, imbricate.

10.3897/zookeys.1040.65750.figure4C2DFA215-FC7D-573B-B95A-A6FE7F6093E9

Male holotype of Cyrtodactylus macrotuberculatus from Pulau Langkawi, Kedah, Peninsular Malaysia (ZRC 2.6754) in preservative A dorsal view B ventral view C tuberculate gular region and throat, and D enlarge femoroprecloacal scale row and pores.

https://binary.pensoft.net/fig/548920
Coloration of adult male <abbrev xmlns:xlink="http://www.w3.org/1999/xlink" content-type="institution" xlink:title="Zoological Museum, Kasetsart University, Thailand" id="ABBRID0ERGDK" xlink:type="simple">ZMKU</abbrev> R 00871 in life

(Fig. 5). Ground color of head, body, limbs, and dorsum light-brown to yellowish brown; wide, dark-brown nuchal band edged anteriorly and posteriorly by thin, creamy-white lines bearing tubercles extends from posterior margin of one eye to posterior margin of other eye; four similar body bands between nuchal loop and hind limb insertions edged anteriorly and posteriorly by thin, creamy-white lines bearing tubercles, first band terminates at shoulders, second and third bands terminate just dorsal to ventrolateral folds, the fourth band terminates at femurs; dark body bands slightly larger than light-colored interspaces; one additional dark-brown band posterior to hind limbs; original portion of tail bearing eight ringed, dark-colored bands separated by seven, narrower, off-white bands infused with dark pigmentation; ventral surfaces of head smudged with brown; abdomen and limbs beige, slightly darker, lateral regions.

10.3897/zookeys.1040.65750.figure546662B51-F31D-5EF8-B27B-2C13547AB5F9

Adult male Cyrtodactylus macrotuberculatus from Adang Island, Satun Province, Thailand (ZMKU R 00871) in life.

https://binary.pensoft.net/fig/548921
Coloration in preservative

(Fig. 6). Color pattern of head, body, limbs, and tail similar to that in life with some fading. Ground color of head, body, limbs, and dorsum tan; dark body and dark caudal bands lighter than in life.

10.3897/zookeys.1040.65750.figure68CC7CAB5-A60D-58E1-A919-376B03A777C6

Adult male Cyrtodactylus macrotuberculatus from Adang Island, Satun Province, Thailand (ZMKU R 00871) in preservative A dorsal and B ventral views.

https://binary.pensoft.net/fig/548922
Variation.

Cyrtodactylus macrotuberculatus usually varies in coloration and banding pattern (Figs 78). In females, a precloacal groove and pores are absent (Fig. 9). PSUZC-RT 2010.58 and THNHM 15378 have a shallow precloacal groove. Three dark body bands occur in PSU 2010.58, THNHM 15378, ZMKU R 00889–00894 and ZMKU R 00897. In ZMKU R 00887, the second dorsal band bifurcates just dorsal to the ventrolateral fold. ZMKU R 00895 has four bands and the third band is incomplete. The third body band in ZMKU R 00896 is broken on the left of the midline and contacts the fourth body band bilaterally. Nuchal loop and body bands of ZMKU R 00883, ZMKU R 00895, and ZMKU R 00898 edged anteriorly and posteriorly by thin, light-yellow lines and tubercles; and dorsal superciliaries are light-yellow (Fig. 8). Variation in morphometric and meristic data are shown in Table 6.

10.3897/zookeys.1040.65750.figure7A1853A3E-0891-5BB6-8A1D-4F5CD5A16B88

Variation in dorsal body band pattern of Cyrtodactylus macrotuberculatus from Thailand. From left to right, upper: ZMKU R 00878, ZMKU R 00873 from Adang Island, Satun Province; and ZMKU R 00887 from Rawi Island, Satun Province. Lower: ZMKU R 00889 from Rawi Island, Satun Province; ZMKU R 00896 and ZMKU R 00895 from Phuket Province.

https://binary.pensoft.net/fig/548923
10.3897/zookeys.1040.65750.figure826BD5EAA-5FA4-5197-BABF-80A56A7217A6

Color in life of Cyrtodactylus macrotuberculatus from Thailand A adult male ZMKU R 00883 from Rawi Island, Satun Province B subadult female ZMKU R 00895 from Thalang District, Phuket Province, and C juvenile ZMKU R 00898 from Kathu District, Phuket Province.

https://binary.pensoft.net/fig/548924
10.3897/zookeys.1040.65750.figure905247188-4F85-52DB-BBBE-F645361E15B2

Precloacal region in female specimens of Cyrtodactylus macrotuberculatusA paratype ZRC 2.6758, from Telaga Tujuh, Pulau Langkawi, Malaysia, and BZMKU R 00896 from Thalang District, Phuket Province, Thailand.

https://binary.pensoft.net/fig/548925
Distribution.

Cyrtodactylus macrotuberculatus is distributed on the mainland and only known from one island in Peninsular Malaysia and southern Thailand (Fig. 1). This species is known from Pulau Langkawi (Gunung Raya, Telaga Tujuh, Gunung Machinchang, and Lubuk Semilang), Kedah, Peninsular Malaysia (Grismer and Ahmad 2008). Other populations are found from Peninsular Malaysia; Kedah (Bukit Wang, Gunung Jerai, Hutan Lipur Sungai Tupah, Kuala Nerang, and Ulu Muda) and Perlis (Bukit Chabang, Chuping and Perlis State Park; [Grismer 2011; Grismer et al. 2012; Quah et al. 2019]). In Thailand, C. macrotuberculatus was recorded from Phatthalung Province (Grismer et al. 2012); Phuket Province, Kathu District (Kathu Waterfall) and Thalang District (Thep Krasatti Sub-district, previously type locality of C. phuketensis); Satun Province, La-ngu District, and Mueang Satun District (Adang and Rawi Islands); Songkhla Province, Rattaphum District (Grismer et al. 2012) and Hat Yai District (Ton Nga Chang Waterfall).

Natural history.

Based on specimens in Thailand, all individuals were found in similar habitat type, lowland forest habitat along granitic rock streams and surrounding areas (elevation 7–186 m asl) during a night survey (1900–2200; Fig. 10). The geckos were found mostly on rock boulders, vegetation (trunk of tree, buttress root, rotting wood and vines), and sometimes on the ground with leaf litter and high humidity (26.3–30.8 °C in temperature, 73.8–100% in relative humidity). Gravid female (ZMKU R 00876) contained four eggs during December. One juvenile (ZMKU R 00898, 56.50 mm in SVL) was found on a tree trunk in January. The varied microhabitats within which this species occurs, are consistent with its characterization as a habitat generalist (Grismer et al. 2020, 2021b) and may account for its wide peninsular and insular distribution relative to other species of the pulchellus group whose distributions are much less extensive or site-specific (Grismer et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020).

10.3897/zookeys.1040.65750.figure10BC6F9A34-62A1-5A5D-BAAD-639FF3597D50

Habitats of Cyrtodactylus macrotuberculatus in Thailand A Adang Island, Satun Province B Kathu District, Phuket Province, and C Hat Yai District, Songkhla Province.

https://binary.pensoft.net/fig/548926

In Thailand, C. macrotuberculatus were found sympatric with other gecko species, Cnemaspis adangrawiAmpai et al., 2019 on Adang and Rawi Islands, Satun Province (Ampai et al. 2019); Cnemaspis phuketensis Das and Leong, 2004, Cyrtodactylus oldhami Theobald, 1876, and Gekko (Ptychozoon) tokehos Grismer et al., 2019 at Kathu and Thalang District, Phuket Province; G. (P.) tokehos, Cnemaspis kumpoli Taylor, 1963, and Gehyra mutilata (Weigmann, 1834) at Hat Yai District, Songkhla Province.

Comparison.

Cyrtodactylus macrotuberculatus is distinguished from all other 15 species in the C. pulchellus complex by a combination of morphological characters (Table 7). It differs from all other species by having prominent tuberculation on the body; tubercles on ventral surface of forelimbs, gular region, and in ventrolateral body folds; 34–49 paravertebral tubercles; 19–27 longitudinal tubercle rows; 17–28 ventral scales; 19–23 subdigital lamellae on the fourth toe; 28–42 femorprecloacal pores in males; deep precloacal groove in males; no scattered white spots on dorsum; 7–10 dark-ringed caudal bands on original tail; white caudal bands on original tail infused with dark pigmentation in adults. Additional comparisons between C. macrotuberculatus and other species in C. pulchellus complex are in Table 7.

Diagnostic characters of Cyrtodactylus macrotuberculatus and related species within C. pulchellus complex. W = weak; P = prominent; / = data unavailable. Some information was collected from the following literature (Grismer et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020).

australotitiwangsaensis bintangtinggi bintangrendah evanquahi lenggongensis pulchellus sharkari trilatofasciatus macrotuberculatus (this study) astrum dayangbuntingensis hidupselamanya langkawiensis lekaguli jelawangensis timur
Supralabials9–129–138–129 or 1010 or 119–111110–129–1310–1212–149–129–1210–129–1210–12
Infralabials9–138–118 or 109 or 108–108–10108–117–119–1210–118–118–109–119–118–10
TuberculationPPPPWPWPPWWWWWPW
Tubercles on ventral surface of forelimbsNoNoNoNoNoNoNoNoYesNoNoNoNoNoYesNo
Tubercles in gular regionNoNoNoNoNoNoNoNoYesNoNoNoNoNoNoNo
Ventrolateral fold tuberculateNoNoYesNoNoNoNoNoYesNoNoNoNoNoNoNo
Paravertebral tubercles37–4531–4236–4431–3436–4133–433134–3834–4940–5735–3639–4834–4430–5036–4238–43
Longitudinal rows of tubercles22–3021–2622–2518–2322–2522–262423–2719–2720–2920–2219–2421–2520–2423–2521–24
Ventral scales32–4036–4031–3929–3332 or 3329–344133–3617–2831–4636–3926–3338–4331–4331–3631–40
4th toe lamellae21–2521–2421–2422–2320–2321–262422–2719–2320–2421–2319–2419–2120–2521–2421–25
Femoroprecloacal pores39–4537–4141–4632–3639–4133–394641–4628–4231–3826–2917–223030–363621 or 22
Precloacal groove in malesDeepDeepDeepShallowDeepDeepShallowDeepDeepDeepDeepDeepDeepDeepDeepDeep
No. of body bands3(1) or 43(1) or 446 or 74 or 54433–44444 or 54 or 544
Body band/interspace ratio1.00–2.001.00–1.251.00–1.250.82–1.100.50–1.250.75–1.251.752.00–2.750.95–1.741.00–2.000.751.00–1.250.75–1.001.00–2.001.00–1.501.00–1.25
Dorsum bearing scattered pattern of white tuberclesNoNoNoNoNoNoNoNoNoYesYesNoNoNoNoNo
Hatchlings/juveniles with white tail tipNoNoNoYes/No/NoNoYesYesYesYesYesYesNo
Adult posterior caudal region whiteNoNoNoYesNoNoNoNoNoNoNoYesNoNoNoNo
Dark caudal bands on original tail7–88–108 or 99–11148–1076–77–1013 or 14>78–1011–1612–14108–10
White caudal bands in adults immaculateYesYesYesNoYesYesYesYesNoNoNoYesNoNoNoYes
Maximum SVL120.10111.1114.4096.00103.1114.1100.1122.2117.9108.399.00102.799.8103.5119.8120.5
Sample size121463413163911314101345

Based on molecular data, C. macrotuberculatus is the sister lineage to a clade composed of C. pulchellus and C. evanquahi. It can be separated from those two species by having tubercles on ventral surface of forelimbs, gular region, and in ventrolateral body folds (vs. absent in C. evanquahi and C. pulchellus); 17–28 ventral scales (vs. 29–33 in C. evanquahi and 29–34 in C. pulchellus); deep precloacal groove in males (vs. a shallow in C. evanquahi); three or four dark dorsal bands (vs. six or seven bands in C. evanquahi and only four bands in C. pulchellus); white posterior caudal region absent (vs. present in C. evanquahi); hatchlings and juveniles without white tail tip (vs. present in C. evanquahi).

Specimens of Cyrtodactylus used in (A) molecular and/or (B) morphological analyses in this study. WM = West Malaysia; TH = Thailand.

SpeciesLocalityMuseum No.GenBank Accession No.Type of analysisReference
Hemidactylus frenatus UnknowNC 00155 JX519468 A Grismer et al. (2012)
Agamura persica Pakistan, Baluchistan Province, Makran District, GwadarFMNH 247474 JX440515 A Grismer et al. (2012)
Tropiocolotes steudneri UnknowJB 28 JX440520 A Grismer et al. (2012)
Cyrtodactylus elok WM, Pahang, Fraser’s Hill, The GapLSUHC 6471 JQ889180 A Grismer et al. (2012)
C. intermedius TH, Chantaburi Province, Khao Khitchakut DistrictLSUHC 9513 JX519469 A Grismer et al. (2012)
TH, Chantaburi Province, Khao Khitchakut DistrictLSUHC 9514 JX519470 A Grismer et al. (2012)
Laos, Khammouan Province, Nakai DistrictFMNH 255454 JQ889181 A Grismer et al. (2012)
Cyrtodactylus sp.TH, Loei, Phu RuaFMNH 265806 JX519471 A Grismer et al. (2012)
C. astrum WM, Perlis, Gua KelamLSUHC 8806 JX519481 A Grismer et al. (2012)
WM, Perlis, Gua KelamLSUHC 8808 JX519479 A Grismer et al. (2012)
WM, Perlis, Kuala PerlisLSUHC 8815 JX519482 A Grismer et al. (2012)
WM, Perlis, Kuala PerlisLSUHC 8816 (paratype) JX519483 A Grismer et al. (2012)
C. australotitiwangsaensis WM, Pahang, Genting HighlandsLSUHC 6637 (holotype) JX519484 A Grismer et al. (2012)
WM, Pahang, Fraser’s HillLSUHC 8086 JX519486 A Grismer et al. (2012)
WM, Pahang, Fraser’s HillLSUHC 8087 JX519485 A Grismer et al. (2012)
C. bintangrendah WM, Kedah, Bukit PalangLSUHC 9984 JX519487 A Grismer et al. (2012)
WM, Kedah, Bukit MertajamLSUHC 10331 (paratype) MN125076 A Quah et al. (2019)
WM, Kedah, Bukit MertajamLSUHC 10519 MN125077 A Quah et al. (2019)
WM, Kedah, Bukit MertajamLSUHC 10520 (paratype) MN125078 A Quah et al. (2019)
C. bintangtinggi WM, Perak, Bukit LarutLSUHC 8862 JX519493 A Grismer et al. (2012)
WM, Perak, Bukit LarutLSUHC 9006 (paratype) JX519494 A Grismer et al. (2012)
C. dayangbuntingensis WM, Kedah, Dayang Bunting IslandLSUHC 14353 MN125090 A Quah et al. (2019)
WM, Kedah, Dayang Bunting IslandLSUHC 14354 MN125091 A Quah et al. (2019)
WM, Kedah, Dayang Bunting IslandLSUHC 14355 MN125092 A Quah et al. (2019)
C. evanquahi WM, Kedah, Gunung BalingBYU 53435 (holotype) MN586889 A Wood et al. (2020)
WM, Kedah, Gunung BalingBYU 53436 (paratype) MN586890 A Wood et al. (2020)
WM, Kedah, Gunung BalingBYU 53437 (paratype) MN586891 A Wood et al. (2020)
C. hidupselamanya WM, Kelantan, Felda Chiku 7LSUHC 12158 (paratype) KX011412 A Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7LSUHC 12160 (paratype) KX011414 A Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7LSUHC 12161 (paratype) KX011415 A Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7LSUHC 12162 (paratype) KX011416 A Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7LSUHC 12163 (holotype) KX011417 A Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7LSUHC 12173 (paratype) KX011420 A Grismer et al. (2016)
C. jelawangensis WM, Gunung Stong, KelantanLSUHC 11060 (paratype) KJ659850 A Grismer et al. (2014)
WM, Kelantan, Gunung StongLSUHC 11061 (paratype) KJ659851 A Grismer et al. (2014)
WM, Gunung Stong, KelantanLSUHC 11062 (holotype) KJ659852 A Grismer et al. (2014)
C. langkawiensis WM, Kedah, Pulau Langkawi, Wat WanaramLSUHC 9123 (paratype) JX519500 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat WanaramLSUHC 9124 (paratype) JX519499 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat WanaramLSUHC 9125 JX519496 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat WanaramLSUHC 9435 JX519495 A Grismer et al. (2012)
C. lekaguli TH, Phang-nga Province, Takua Pa DistrictZMKU 00720 KX011425 A Grismer et al. (2016)
TH, Phang-nga Province, Takua Pa DistrictZMKU 00721 KX011426 A Grismer et al. (2016)
TH, Phang-nga Province, Takua Pa DistrictZMKU 00722 KX011427 A Grismer et al. (2016)
TH, Phang-nga Province, Takua Pa DistrictZMKU 00723 KX011428 A Grismer et al. (2016)
C. lenggongensis WM, Perak, Lenggong ValleyLSUHC 9974 (holotype) JX519490 A Grismer et al. (2012)
WM, Perak, Lenggong ValleyLSUHC 9975 (paratype) JX519488 A Grismer et al. (2012)
WM, Perak, Lenggong ValleyLSUHC 9976 (paratype) JX519489 A Grismer et al. (2012)
WM, Perak, Lenggong ValleyLSUHC 9977 (paratype) JX519491 A Grismer et al. (2012)
C. macrotuberculatus WM, Kedah, Kuala NerangBYU 51869 MN125085 A Quah et al. (2019)
WM, Kedah, Kuala NerangBYU 51870 MN125086 A Quah et al. (2019)
WM, Kedah, Gunung JeraiLSUHC 5939 JX519513 A Grismer et al. (2012)
WM, Kedah, Gunung JeraiLSUHC 5999 JX519512 A Grismer et al. (2012)
WM, Kedah, Gunung JeraiLSUHC 6000 JX519514 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Lubuk SembilangLSUHC 6829 JX519505 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Gunung MachinchangLSUHC 7560 JX519503 A Grismer et al. (2012)
C. macrotuberculatus WM, Kedah, Pulau Langkawi, Gunung RayaLSUHC 9428 JX519506 A, BGrismer et al. (2012), This study
WM, Kedah, Pulau Langkawi, Gunung RayaLSUHC 9429BThis study
WM, Kedah, Pulau Langkawi, Gunung RayaLSUHC 9432BThis study
WM, Kedah, Pulau Langkawi, Gunung MachinchangLSUHC 9448 JX519507 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Gunung MachinchangLSUHC 9449 JX519509 A Grismer et al. (2012)
WM, Kedah, Hutan Lipur Sungai TupahLSUHC 9671 JX519510 A Grismer et al. (2012)
WM, Kedah, Hutan Lipur Sungai TupahLSUHC 9672 JX519511 A Grismer et al. (2012)
WM, Kedah, Hutan Lipur Sungai TupahLSUHC 9693 JX519517 A Grismer et al. (2012)
WM, Perlis, Perlis State ParkLSUHC 9980 JX519515 A Grismer et al. (2012)
WM, Perlis, Perlis State ParkLSUHC 9981 JX519516 A, BGrismer et al. (2012), This study
WM, Perlis, Bukit ChabangLSUHC 10037 JX519519 A Grismer et al. (2012)
WM, Perlis, Bukit ChabangLSUHC 10038 JX519518 A Grismer et al. (2012)
WM, Perlis, Perlis State ParkLSUHC 10067BThis study
WM, Kedah, Bukit WangLSUHC 10329 MN125088 A Quah et al. (2019)
WM, Kedah, Bukit WangLSUHC 10330 MN125087 A Quah et al. (2019)
WM, Perlis, Perlis State ParkZRC 2.4869BThis study
WM, Kedah, Pulau Langkawi, Gunung RayaZRC 2.6754 (holotype)BThis study
WM, Kedah, Pulau Langkawi, Gunung RayaZRC 2.6755 (paratype)BThis study
WM, Kedah, Pulau Langkawi, Gunung RayaZRC 2.6756 (paratype)BThis study
WM, Kedah, Pulau Langkawi, Telaga TujuhZRC 2.6757/ LSUHC 7173 (paratype) JX519508 A Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Lubuk SemilangZRC 2.6758 (paratype)BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00871BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00872BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00873BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00874 MW809294 A, BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00875 MW809295 A, BThis study
TH, Songkhla Province, Hat Yai District, Thung Tam SaoZMKU R 00876 MW809296 A, BThis study
TH, Songkhla Province, Hat Yai District, Thung Tam SaoZMKU R 00877 MW809297 A, BThis study
TH, Songkhla Province, Hat Yai District, Thung Tam SaoZMKU R 00878 MW809298 A, BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00879BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00880BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00881BThis study
TH, Satun Province, Mueang Satun District, Adang IslandZMKU R 00882BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00883 MW809299 A, BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00884BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00885BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00886BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00887 MW809300 A, BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00888BThis study
TH, Satun Province, Mueang Satun District, Rawi IslandZMKU R 00889BThis study
C. macrotuberculatus (as C. phuketensis)TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00890 MW809301 A, BThis study
TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00891 MW809302 A, BThis study
TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00892 MW809303 A, BThis study
TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00893 MW809304 A, BThis study
TH, Phuket Province, Thalang District, Thep KrasattiZMKU R 00894 MW809305 A, BThis study
TH, Phuket Province, Thalang District, Thep KrasattiZMKU R 00895 MW809306 A, BThis study
TH, Phuket Province, Thalang District, Thep KrasattiZMKU R 00896 MW809307 A, BThis study
TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00897 MW809308 A, BThis study
TH, Phuket Province, Kathu District, Kathu WaterfallZMKU R 00898 MW809309 AThis study
TH, Phuket Province, Thalang District, Thep KrasattiPSUZC-RT 2010.58BThis study
TH, Phuket Province, Thalang District, Thep KrasattiTHNHM 15378BThis study
C. pulchellus WM, Penang, Pulau Pinang, Empangan Air ItamLSUHC 6668 JX519523 A Grismer et al. (2012)
WM, Penang, Pulau Pinang, Moongate TrailLSUHC 6726 JX519527 A Grismer et al. (2012)
WM, Penang, Pulau Pinang, Moongate TrailLSUHC 6727 JX519526 A, BGrismer et al. (2012), This study
WM, Penang, Pulau Pinang, Moongate TrailLSUHC 6728 JX519525 A, BGrismer et al. (2012), This study
WM, Penang, Pulau Pinang, Moongate TrailLSUHC 6729 JX519528 A, BGrismer et al. (2012), This study
WM, Penang, Pulau Pinang, Moongate TrailLSUHC 6785 JX519524 A Grismer et al. (2012)
WM, Penang, Pulau Pinang, Air Terjun Titi KerawangLSUHC 9667 JX519520 A Grismer et al. (2012)
WM, Penang, Pulau Pinang, Air Terjun Titi KerawangLSUHC 9668 JX519521 A Grismer et al. (2012)
WM, Penang, Pulau Pinang, Air Terjun Titi KerawangLSUHC 10022 JX519522 A, BGrismer et al. (2012), This study
WM, Penang, Pulau PinangZRC 2.4854BThis study
WM, Penang, Pulau PinangZRC 2.4857BThis study
C. sharkari WM, Pahang, Merapoh, Gua GuntingLSUHC 11022 (holotype) KJ659853 A Grismer et al. (2014)
C. timur WM, Gunung Tebu, TerengganuLSUHC 10886 KJ659854 A Grismer et al. (2014)
WM, Gunung Tebu, TerengganuLSUHC 11183 (paratype) KJ659855 A Grismer et al. (2014)
WM, Gunung Tebu, TerengganuLSUHC 11184 (paratype) KJ659856 A Grismer et al. (2014)
WM, Gunung Tebu, TerengganuLSUHC 11185 (paratype) KJ659857 A Grismer et al. (2014)
C. trilatofasciatus WM, Pahang, Cameron HighlandsLSUHC 10064 JX519529 A Grismer et al. (2012)
WM, Pahang, Cameron HighlandsLSUHC 10065 JX519530 A Grismer et al. (2012)
WM, Pahang, Cameron HighlandsLSUHC 10066 JX519531 A Grismer et al. (2012)
10.3897/zookeys.1040.65750.figure10E3FBA02-E635-5675-961A-681911D81FFA

Map illustrating the known geographic distribution of Cyrtodactylus macrotuberculatus and C. phuketensis. Yellow star: the type locality of C. macrotuberculatus at Gunung Raya, Pulau Langkawi, Kedah, Malaysia. Green star: the type locality of C. phuketensis at Thalang District, Phuket Island, Phuket Province. Yellow circles: C. macrotuberculatus samples used in this study. Green circle: C. phuketensis samples used in this study. Yellow squares: the distribution of C. macrotuberculatus taken from Grismer et al. (2012), and Quah et al. (2019). The samples used correspond to those in Table 1.

https://binary.pensoft.net/fig/548917
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