Animalia Squamata Gekkonidae TermprayoonKorkhwanRujirawanAttapolGrismerL. LeeWood JrPerry L.AowpholAnchaleeTwo new karst-adapted species in the Cyrtodactyluspulchellus group (Reptilia, Gekkonidae) from southern ThailandZookeys14092023117931335210.3897/zookeys.1179.109712 F7410CEC-A0A3-54B4-84B2-06C704D5D478 Cyrtodactylus wangkhramensis https://zoobank.org/D2615DAF-F982-41C0-85A1-4C8D93816E99 sp. nov.Figs 9, 10, 11 Wangkhram Bent-toed GeckoType material.

Holotype. Adult male (ZMKU R 01018, Figs 9, 10) collected from Thailand, Satun Province, La-ngu District, Khao Khao Subdistrict, Tham (= cave) Wangkhram (6°56.324'N, 99°48.920'E; 0 m a.s.l.), on 13 March 2019 by Korkhwan Termprayoon, Anchalee Aowphol, Attapol Rujirawan, Natee Ampai and Siriporn Yodthong.

10.3897/zookeys.1179.109712.figure9B6FC4B47-41CE-53B3-9B64-14E8F5F52100

Adult male holotype of Cyrtodactyluswangkhramensis sp. nov. (ZMKU R 01018) from the type locality at Tham Wangkhram, La-ngu District, Satun Province, Thailand A dorsolateral view of specimen in life B dorsal and ventral views immediately after euthanasia.

https://binary.pensoft.net/fig/905282
10.3897/zookeys.1179.109712.figure10EB28BB30-44C0-5694-AED7-95BE7AD82F69

Adult male holotype of Cyrtodactyluswangkhramensis sp. nov. (ZMKU R 01018) in preservation A dorsal and ventral views of body B head dimensions showing lateral, dorsal and ventral views C ventrolateral fold on the left side D precloacal depression and pore-bearing femoroprecloacal scales E dorsal view of tail showing last row tubercles.

https://binary.pensoft.net/fig/905283

Paratypes. Eleven paratypes, four adult males (ZMKU R 01015–01017 and ZMKU R 01019) and seven adult females (ZMKU R 01020–01026), same data as holotype.

Diagnosis.

Cyrtodactyluswangkhramensis sp. nov. can be distinguished from all other species of the C.pulchellus group by a combination of the following characters: (1) SVL 87.4–95.5 mm in adult males (n = 5), 89.4–98.8 mm in adult females (n = 7); (2) 11–14 supralabial and 9–13 infralabial scales; (3) weak tuberculation on body; (4) no tubercles on ventral surfaces of forelimbs, gular region or in ventrolateral body folds; (5) 28–35 paravertebral tubercles; (6) 19–21 longitudinal rows of dorsal tubercles; (7) 34–40 rows of ventral scales; (8) 18–22 subdigital lamellae on the fourth toe; (9) 32–36 femoroprecloacal pores in adult males; (10) absence of precloacal pores in adult females; (11) deep precloacal groove in males; (12) absence of scattered pattern of white tubercles on dorsum; (13) four or five dark dorsal body bands; (14) 11 dark caudal bands on original tail; (15) light caudal bands in adults infused with dark pigmentation; and (16) caudal tubercles extending 1/5–1/7 of anterior portion of tail.

Description of holotype.

Adult male SVL; 94.2 mm; head moderate in length (HL/SVL 0.28) and wide (HW/HL 0.63), flattened (HD/HL 0.38), distinct from neck and triangular in dorsal profile; lores concave anteriorly, inflated posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis rounded anteriorly; snout elongate (ES/HL 0.41), rounded in dorsal profile, laterally constricted; eye large (ED/HL 0.23); ear opening elliptical, moderate in size (EL/HL 0.08), obliquely orientated; eye to ear distance slightly greater than diameter of eye; rostral rectangular, divided dorsally by an inverted U-shaped furrow, bordered posteriorly by left and right supranasals and large hexagonal internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by a large anterior supranasal, posteriorly by two postnasals, ventrally by first supralabial; 8/8 (left/right) rectangular supralabials extending to below mid-point of eye, 12/12 to below the posterior margin of the eye-ball, decreasing abruptly just posterior to mid-point of eye; 7/7 infralabials extending to below mid-point of eye, 11/11 to upturn the labial margin, decreasing gradually 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 rounded, small tubercles; dorsal superciliaries elongate, smooth, largest anteriorly; mental triangular, 3.60 mm in width, 2.38 mm in length, bordered laterally by first infralabials and posteriorly by left and right, trapezoidal postmentals which contact medially for approximately 45% of their length; one row of slightly enlarged, elongate sublabials extending posteriorly to the seventh (left/right) infralabials; small, granular, gular scales grading posteriorly into larger, flat, smooth, imbricate, pectoral and ventral scales.

Body relatively short (AG/SVL 0.47) with well-defined, non-tuberculate, ventrolateral folds; dorsal scales small, granular, interspersed with low, regularly arranged, weakly-keeled tubercles, smaller intervening tubercles occasionally present; tubercles extending from occiput to base of tail, but not further than 1/5 of tail; tubercles on occiput and nape relatively small, those on body largest; approximately 20 longitudinal rows of tubercles at mid-body; 32 paravertebral tubercles; 35 flat imbricate ventral scales between ventrolateral body folds; ventral scales larger than dorsal scales; precloacal scales large, smooth; deep precloacal groove.

Forelimbs moderately slender, relatively short (FL/SVL 0.17); dorsal scales on forelimbs slightly raised, granular, slightly larger than those on body intermixed with enlarged, subconical and weakly-keeled tubercles; scales of ventral surface of forearm flat, subimbricate, tubercles absent; palmar scales small, weakly rounded; digits well-developed, inflected at basal, interphalangeal joints; 17/17 (left/right) subdigital lamellae on the fourth finger, 6/6 proximal subdigital lamellae rectangular, broadly expanded proximal to joint inflection, 11/11 distal subdigital lamellae slightly expanded distal to inflection becoming gradually more expanded near the claw; claws well-developed, sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.20), enlarged, subconical, weakly-keeled tubercles on dorsal surface of legs separated by smaller juxtaposed scales; ventral scales of thigh flat, smooth, imbricate, larger than dorsal granular scales; ventral, tibial scales flat, smooth, imbricate; a single row of 34 enlarged femoroprecloacal scales extending nearly from knee to knee through precloacal region where they are continuous with enlarged, pore-bearing precloacal scales; 32 contiguous pore-bearing femoroprecloacal scales (Fig. 10D), forming an inverted T bearing a deep, precloacal groove; seven pore-bearing scales bordering groove (four on left and three on right); postfemoral scales immediately posterior to enlarged scale row small, nearly granular, forming an abrupt union with postfemoral scales on posteroventral margin of thigh; plantar scales weakly rounded to flat; 18/18 subdigital lamellae on fourth toe, 7/7 proximal subdigital lamellae rectangular, broadly expanded proximal to joint inflection, 11/11 distal subdigital lamellae slightly expanded distal to inflection becoming gradually more expanded near the claw; claws well-developed, sheathed by dorsal and ventral scales.

Regenerated tail 120.3 mm in length, slightly longer than SVL (TL/SVL = 1.28), 5.4 mm in width at base, tapering to a point; dorsal scales of tail flat, squarish; original portion segmented, 7–9 transverse scales rows per segment; seven transverse rows of 2–6 dorsal tubercles on posterior margin, fewer posteriorly; subcaudal region bearing large median row of transverse scales; regenerated portion of tail covered with small, smooth, rectangular scales dorsally and ventrally; shallow dorsal and lateral caudal furrow extending entire length of original tail; base of tail bearing hemipenial swellings; two rows of (3+2)L/(1+3)R medium-sized postcloacal tubercles on each hemipenial swelling; postcloacal scales smooth, flat, large, imbricate.

Colouration in life

(Fig. 9). Ground colour of body, limbs, light-brown, those on head lighter; faded, incomplete V-shaped rostral chevron; superciliaries yellow; supralabial and infralabial scales light-brown anteriorly, with off-white-coloured medially to posteriorly; wide, dark-brown nuchal band edged anteriorly and posteriorly by thin, white lines bearing tubercles extending from posterior margin of one eye to posterior margin of another eye; four dark-brown body bands between nuchal loop and hind limb insertions edged anteriorly and posteriorly by broken, thin, white lines formed by a single row of white tubercles, first band terminating at shoulders, second and third bands terminating just dorsal to ventrolateral folds, the fourth band terminating at femurs; dark body bands slightly larger than light-coloured interspaces; creamy pale yellow tubercles on dorsal surfaces of limbs, those on body darker, similar to ground colour; one additional dark-brown band posterior to hind limbs (postsacral band); ventral surfaces of head, abdomen and limbs greyish-white; creamy pale yellow postcloacal tubercles; tail bearing approximately six dark bands separated by six light-brown (anteriorly) to white (posteriorly) bands, white caudal band heavily infused with dark-brown pigmentation on original portion; regenerated portion and subcaudal region tan.

Colouration in preservative

(Fig. 10). The overall colour pattern of head, body, limbs and tail similar to that in life with some fading. Ground colour of head, body, limbs and dorsum dark tan; dark body bands darker; creamy pale yellow-coloured tuberculation on dorsum fading to off-white; beige coloured on the ventral surface.

Variation.

Meristic and morphometric data for the type series of Cyrtodactyluswangkhramensis sp. nov. are given in Table 8 and Suppl. material 3. All paratypes resemble the holotype in general aspects of morphology with variations in colouration and banding pattern. Eight specimens have four dark body bands and three specimens (ZMKU R 01016–01017 and ZMKU R 01021) have five dark bands on dorsum (Fig. 11A). Female paratype (ZMKU R 01025) has four dark body bands with an irregular pattern on the third (Fig. 11B). Original tails (ZMKU R 01019–01021) have 11 dark caudal bands and 10–11 light caudal bands. Last row of caudal tubercles of paratypes (ZMKU R 01019–01021) extending to 4th–7th segment of original portion, approximately 1/5–1/7 of anterior portion of original tail.

Descriptive measurements (millimetres), meristic (left/right) and non-meristic characters of the type series of Cyrtodactyluswangkhramensis sp. nov. Key: H = holotype, P = paratype, M = male, F = female; Or = original tail, Br = broken, Re = regenerated; / = data unavailable or inapplicable; L = left, R = right. Abbreviations are defined in Materials and methods.

ZMKU R 01018ZMKU R 01015ZMKU R 01016ZMKU R 01017ZMKU R 01019ZMKU R 01020ZMKU R 01021ZMKU R 01022ZMKU R 01023ZMKU R 01024ZMKU R 01025ZMKU R 01026
Type seriesHPPPPPPPPPPP
SexMMMMMFFFFFFF
SVL 94.289.795.592.587.493.495.298.892.395.993.789.4
Tail conditionReReReBrOrOrOrReReBrReRe
TL 120.3122.591.8/119.5126.6120.493.9116.551.3111.288.3
TW 5.47.46.6/5.45.25.86.15.65.85.85.0
FL 15.615.515.215.714.815.316.216.415.815.515.315.0
TBL 18.518.518.717.718.117.418.118.818.618.718.417.2
AG 44.645.545.144.841.147.147.049.245.147.145.143.1
HL 26.426.827.527.525.826.827.528.827.828.627.926.4
HW 16.717.517.517.616.117.217.818.317.817.517.316.5
HD 10.011.510.610.59.810.110.710.810.110.210.79.9
ED 6.26.77.06.76.36.16.76.56.76.56.36.1
EE 7.07.17.17.06.96.77.07.06.86.56.96.7
ES 10.711.111.310.410.310.211.211.510.911.010.910.2
EN 8.28.58.68.18.07.88.58.98.38.48.38.0
IO 5.76.26.26.55.45.96.46.45.66.06.16.2
EL 2.02.11.61.92.11.92.32.81.92.02.12.2
IN 3.03.32.72.82.73.13.43.53.43.33.13.0
SL 12/1213/1213/1313/1312/1211/1313/1312/1312/1213/1212/1413/13
SL-mid-eye 8/87/710/99/99/97/99/99/99/99/99/1010/10
IL 11/1111/1111/1311/1111/1111/1112/1011/1111/1111/1111/119/11
IL-mid-eye 7/79/97/88/87/87/79/79/77/88/87/87/8
PVT 323535323334323132352834
LRT 202121212021202119212019
VS 354034353735373634373635
4FLU 11/1111/1213/1311/1111/1212/1213/1212/1213/1313/1312/1212/12
4FLE 6/66/66/66/66/66/66/66/66/66/66/66/6
4FL 17/1717/1819/1917/1717/1818/1819/1818/1819/1919/1918/1818/18
4TLU 11/1112/1214/1413/1212/1213/1313/1313/1313/1314/1312/1213/13
4TLE 7/77/87/78/87/78/88/88/88/88/77/78/8
4TL 18/1819/2021/2121/2019/1921/2121/2121/2121/2122/2019/1921/21
FPP in males3233333632///////
No of pore-bearing scales on precloacal groove7 (4L/3R)6 (3L/3R)6 (3L/3R)6 (3L/3R)7 (3L/4R)///////
PCT rows2L/2R2L/2R2L/1R2L/2R2L/2R///////
No of PCT per row(3+2)L/(1+3)R(2+3)L/(1+4)R(1+4)L/(3)R(3+4)L/(3+4)R(2+4)L/(3+4)R///////
BB 445544544444
LCB ////101110/////
DCB ////111111/////
Body band/ interspace ratio1.701.090.891.311.181.05/0.930.821.251.011.38
Deep precloacal groove in maleYesYesYesYesYes///////
Femoroprecloacal pores continuousYesYesYesYesYes///////
TuberculationWeakWeakWeakWeakWeakWeakWeakWeakWeakWeakWeakWeak
Tubercles on ventral surface of forelimbNoNoNoNoNoNoNoNoNoNoNoNo
Tubercles in gular regionNoNoNoNoNoNoNoNoNoNoNoNo
Ventrolateral fold tuberculateNoNoNoNoNoNoNoNoNoNoNoNo
Dorsum bearing scattered pattern of white tuberclesNoNoNoNoNoNoNoNoNoNoNoNo
Adult posterior caudal region white////NoNoNo/////
White caudal bands in adults immaculate////NoNoNo/////
Portion of caudal tubercles on original tail////1/51/71/7/////
10.3897/zookeys.1179.109712.figure114F39516D-3802-5085-9393-14047A342CE7

Variation in banding pattern of Cyrtodactyluswangkhramensis sp. nov. immediately after euthanasia A adult male (ZMKU R 01016) having five dark body bands on dorsum B adult male (ZMKU R 01026) showing irregular pattern on the 3rd body band.

https://binary.pensoft.net/fig/905284
Distribution.

Cyrtodactyluswangkhramensis sp. nov. is currently known from the type locality at Tham Wangkhram, Khao Khao Subdistrict, La-ngu District, Satun Province, Thailand (Figs 1, 12).

10.3897/zookeys.1179.109712.figure12A76FFA8A-1CE6-5977-9926-9FB8F6F5A916

Habitat of Cyrtodactyluswangkhramensis sp. nov. at the type locality, Tham Wangkhram, La-ngu District, Satun Province, Thailand.

https://binary.pensoft.net/fig/905285
Natural history.

Type series of Cyrtodactyluswangkhramensis sp. nov. was collected from a karst formation at night (1900–2100 h) during March 2019. Eleven specimens were captured from substrates surrounding the karstic area (karst wall, crevices, karst boulders, plantation and concrete poles). Ambient temperature was 25.7 °C and relative humidity was 90.7%. The male holotype (ZMKU R 01018) was found on a termite mound adjacent to a karst outcrop. Five individuals (ZMKU R 01016–01017, ZMKU R 01020–01021 and ZMKU R 01023) were found on a karst wall. One specimen ZMKU R 01015 was found perched on a vine near a karst wall. ZMKU R 01019 was collected from within a karst crevice and ZMKU R 01022 was found on a karst boulder. Two specimens (ZMKU R 01024 and ZMKU R 01026) were perched on concrete poles.

One individual (ZMKU R 01025) was found on a vine in a cave, approximately 10 m from the entrance, where the temperature was 26.1 °C and the relative humidity was 97.9%. Cyrtodactyluswangkhramensis sp. nov. appears to be sympatric with Gehyramutilata (Wiegmann, 1834).

Etymology.

The specific epithet wangkhramensis refers to the type locality at Tham Wangkhram in La-ngu District, Satun Province.

Comparison.

Cyrtodactyluswangkhramensis sp. nov. can be distinguished from other species in the C.pulchellus group by having a combination of weak tuberculation on the body; no tubercles on ventral surface of forelimbs, gular region or in ventrolateral body folds; 11–14 supralabial scales; 28–35 paravertebral tubercles; 19–21 longitudinal tubercle rows; 34–40 ventral scales; 32–36 femorprecloacal pores in males; deep precloacal groove in males; eleven dark caudal bands on original tail; light caudal bands on original tail infused with dark pigmentation in adults; and caudal tubercles 1/5–1/7 of anterior portion of tail. Additional comparisons between Cyrtodactyluswangkhramensis sp. nov. and other species in the C.pulchellus group are in Suppl. material 4.

Cyrtodactyluswangkhramensis sp. nov. is a member of Clade A which comprises Cyrtodactylussungaiupe sp. nov., C.astrum, C.dayangbuntingensis, C.langkawiensis, C.lekaguli and C.stellatus. It differs from those six species by uncorrected pairwise distances of ND2 of 6.59–11.30% (Table 2).

Cyrtodactyluswangkhramensis sp. nov. is sister to Cyrtodactylussungaiupe sp. nov. from which it differs by sequence divergence of 6.59–6.89% (Table 2) and strongly separated by PCA (Fig. 3). Cyrtodactyluswangkhramensis sp. nov. can be differentiated from its sister by having smaller maximum SVL of 98.8 mm (vs. 104.6 mm); caudal tubercles extending between 1/5 and 1/7 of anterior portion of tail (vs. 1/8–1/10 of the tail); and having significantly different mean values of mensural characters of AGadj, HWadj, EEadj and ENadj (p < 0.001–0.009; Table 5). It can be further separated from Cyrtodactylussungaiupe sp. nov. in having statistically significant different mean values of meristic characters of SL (p = 0.045; Table 5).

Cyrtodactyluswangkhramensis sp. nov. differs from C.astrum by having smaller maximum SVL of 98.8 mm (vs. 108.3 mm); 28–35 paravertebral tubercles (vs. 38–57); absence of scattered pattern of white tubercles on dorsum (vs. present); eleven dark caudal bands on the original tail (vs. 13 or 14); caudal tubercles extending between 1/5 and 1/7 of anterior portion of tail (vs. ≥ 1/3 of the tail); and having statistically significant different mean values of mensural characters of SVL, FLadj, TBLadj, HLadj, HWadj, HDadj, EEadj, ESadj, ENadj and ELadj (p < 0.001–0.004; Table 5). It can be further separated from C.astrum in having statistically significant different mean values of meristic characters of PVT and LRT (p < 0.001; Table 5).

Cyrtodactyluswangkhramensis sp. nov. can be separated from C.dayangbuntingensis by having 32–36 femoroprecloacal pores (vs. 26–29); dorsal band interspaces ratio (0.82–1.70 vs. 0.75); absence of scattered pattern of white tubercles on dorsum (vs. present); and one to two rows of postcloacal tubercles (vs. up to three rows).

Cyrtodactyluswangkhramensis sp. nov. differs from C.langkawiensis by having 32–36 femoroprecloacal pores (vs. 30); caudal tubercles extending between 1/5 and 1/7 of anterior portion of tail (vs. ≥ 1/3 of the tail); and having statistically significant different mean values of mensural characters of AGadj, HLadj, ESadj, ENadj and INadj (p = 0.001–0.037; Table 5). Moreover, it can be differentiated from C.langkawiensis in having statistically significant different mean values of meristic characters of SL, IL, PVT, LRT and VS (p < 0.001–0.010; Table 5).

Cyrtodactyluswangkhramensis sp. nov. differs from C.lekaguli by having a smaller maximum SVL of 98.8 mm (vs. 108.5 mm); caudal tubercles extending between 1/5 and 1/7 of anterior portion of tail (vs. ≥ 1/3 of the tail); one to two rows of postcloacal tubercles (vs. one row) and having statistically significant different mean values of mensural characters of SVL, FLadj, HLadj, HWadj, HDadj, EEadj, ESadj, ENadj and IOadj (p < 0.001–0.007; Table 5). It can be differentiated from C.lekaguli in having statistically significant different mean values of meristic characters of SL, PVT and LRT (p = 0.001–0.036; Table 5).

Cyrtodactyluswangkhramensis sp. nov. differs from C.stellatus by having 32–36 femoroprecloacal pores (vs. 24–29); absence of precloacal pores in females (vs. present); absence of scattered pattern of white tubercles on dorsum (vs. present) and having statistically significant different mean values of mensural characters of AGadj, HLadj, HWadj and HDadj (p < 0.001–0.048; Table 5). It can be further separated from C.stellatus in having statistically significant different mean values of meristic characters of PVT (p < 0.001; Table 5).

10.3897/zookeys.1179.109712.suppl3835125632FEB89A-6584-50B5-92FC-E48F7FEAE134

Descriptive morphometric (in millimetres) and meristic characters (min–max) of the type series

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Descriptive morphometric (in millimetres) and meristic characters (min–max) of the type series of Cyrtodactylussungaiupe sp. nov., Cyrtodactyluswangkhramensis sp. nov. and members of Clade A in the Cyrtodactyluspulchellus group. Abbreviations are defined in Materials and Methods. Key: Re = regenerated; L = left; R = right; / = data unavailable or inapplicable.

https://binary.pensoft.net/file/905288This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Korkhwan Termprayoon, Attapol Rujirawan, L. Lee Grismer, Perry L. Wood Jr, Anchalee Aowphol
10.3897/zookeys.1179.109712.figure1BFA5B106-31CA-5C05-9CDD-A46268274732

Map showing the type localities of Cyrtodactylussungaiupe sp. nov. (red star) in Thung Wa District and Cyrtodactyluswangkhramensis sp. nov. (blue star) in La-ngu District, Satun Province, Thailand and the type localities of closely-related species, C.astrum, C.dayangbuntingensis, C.langkawiensis, C.lekaguli and C.stellatus. Circles represent additional localities of specimens used in molecular analyses.

https://binary.pensoft.net/fig/905274
10.3897/zookeys.1179.109712.suppl48351258184338BB-A788-5C86-A080-33C2269300D8

Diagnostic characters of the C.pulchellus group including two new species

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Diagnostic characters of the C.pulchellus group including two new species, Cyrtodactylussungaiupe sp. nov. and Cyrtodactyluswangkhramensis sp. nov. Red shaded cells indicate the differences with Cyrtodactylussungaiupe sp. nov. Blue shaded cells indicate the differences with Cyrtodactyluswangkhramensis sp. nov. Yellow shaded cells indicate the differences with both Cyrtodactylussungaiupe sp. nov. and Cyrtodactyluswangkhramensis sp. nov. Keys: W = weak; P = prominent; / = data unavailable. Additional morphological data were obtained from the following literature (Grismer and Ahmad 2008; Grismer et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020; Termprayoon et al. 2021a, 2021b).

https://binary.pensoft.net/file/905289This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.Korkhwan Termprayoon, Attapol Rujirawan, L. Lee Grismer, Perry L. Wood Jr, Anchalee Aowphol

Percentage uncorrected pairwise sequence divergences (p-distances) of Cyrtodactylussungaiupe sp. nov., Cyrtodactyluswangkhramensis sp. nov. and closely-related species (Clade A), based on 1,444 base pairs of mitochondrial ND2 gene and flanking tRNAs. The p-distance values are given as mean and ranges in parentheses. Intraspecific distances are in bold font.

Species n Cyrtodactylussungaiupe sp. nov.Cyrtodactyluswangkhramensis sp. nov. C.astrum C.dayangbuntingensis C.langkawiensis C.lekaguli C.stellatus
Cyrtodactylussungaiupe sp. nov.7 0.02 (0.00–0.07)
Cyrtodactyluswangkhramensis sp. nov.76.76 (6.59–6.89) 0.28 (0.00–0.52)
C.astrum 129.75 (9.18–10.69)9.63 (8.97–11.01) 1.44 (0.00–3.12)
C.dayangbuntingensis 38.10 (8.02–8.16)7.84 (7.57–8.16)9.56 (9.18–10.90) 0.14 (0.07–0.21)
C.langkawiensis 108.86 (8.67–9.03)8.46 (8.11–8.74)10.06 (9.53–11.54)7.38 (7.17–7.66) 0.28 (0.00–0.52)
C.lekaguli 98.57 (8.11–9.23)8.93 (8.16–9.52)9.74 (8.99–11.59)8.56 (8.00–9.20)9.30 (8.42–10.08) 2.20 (0.00–4.27)
C.stellatus 59.90 (9.59–10.47)10.16 (9.62–11.30)10.54 (9.85–12.34)9.48 (9.21–10.29)10.39 (9.98–11.37)9.22 (8.35–10.60) 0.42 (0.00–0.95)
10.3897/zookeys.1179.109712.figure347363DA1-3889-5C8A-A0C4-5EC672FB79CF

Principal Component Analysis (PCA) of Cyrtodactylussungaiupe sp. nov., Cyrtodactyluswangkhramensis sp. nov. and the closely-related species in Clade A of the C.pulchellus group. The letters in the scatter plots refer to holotype (= H), paratype (= P) and topotype (= T).

https://binary.pensoft.net/fig/905276

Summary pairwise differences of statistically significant characters (Tukey’s test; p < 0.05) from morphometric and meristic characters of Cyrtodactylussungaiupe sp. nov., Cyrtodactyluswangkhramensis sp. nov. and closely-related species (Clade A). Abbreviations are listed in Materials and Methods. Key: * tested by Games-Howell test; ** tested by Dunn’s test. No significantly different characters of 4TL and BB were excluded.

Characters SVL**FLadj* TBL adj AGadj* HL adj HW adj HDadj* EE adj ED adj ESadj**
Cyrtodactylussungaiupe sp. nov. vs. C.astrum< 0.001< 0.0010.006< 0.0010.0160.0040.008
Cyrtodactylussungaiupe sp. nov. vs. C.langkawiensis0.001< 0.0010.033
Cyrtodactylussungaiupe sp. nov. vs. C.lekaguli< 0.0010.0010.008
Cyrtodactylussungaiupe sp. nov. vs. C.stellatus0.0350.0090.002< 0.001< 0.001< 0.001< 0.001< 0.0010.005
Cyrtodactyluswangkhramensis sp. nov. vs. Cyrtodactylussungaiupe sp. nov.0.0030.008< 0.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.astrum0.004< 0.001< 0.001< 0.001< 0.001< 0.001< 0.001< 0.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.langkawiensis0.0280.0370.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.lekaguli< 0.0010.007< 0.001< 0.001< 0.001< 0.001< 0.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.stellatus0.0480.015< 0.0010.014
C.astrum vs. C.langkawiensis0.031< 0.001< 0.001< 0.0010.004< 0.001
C.astrum vs. C.lekaguli< 0.001< 0.0010.0320.030
C.astrum vs. C.stellatus0.002< 0.001< 0.001< 0.001< 0.001< 0.001< 0.001< 0.001
C.langkawiensis vs. C.lekaguli0.0120.005< 0.001< 0.0010.020< 0.001
C.langkawiensis vs. C.stellatus< 0.001< 0.001
C.lekaguli vs. C.stellatus< 0.001< 0.0010.004< 0.001< 0.001< 0.001< 0.001< 0.001< 0.001
Characters ENadj**IOadj* EL adj INadj**SL**IL**PVT**LRT**VS*
Cyrtodactylussungaiupe sp. nov. vs. C.astrum0.0430.0090.0040.011< 0.001< 0.001
Cyrtodactylussungaiupe sp. nov. vs. C.langkawiensis< 0.001< 0.001< 0.0010.046< 0.0010.001
Cyrtodactylussungaiupe sp. nov. vs. C.lekaguli< 0.0010.001< 0.0010.0060.001
Cyrtodactylussungaiupe sp. nov. vs. C.stellatus< 0.0010.003
Cyrtodactyluswangkhramensis sp. nov. vs. Cyrtodactylussungaiupe sp. nov.0.0090.045
Cyrtodactyluswangkhramensis sp. nov. vs. C.astrum< 0.0010.003< 0.001< 0.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.langkawiensis0.0120.0100.0100.0020.005< 0.001< 0.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.lekaguli0.002< 0.0010.0360.0020.001
Cyrtodactyluswangkhramensis sp. nov. vs. C.stellatus< 0.001
C.astrum vs. C.langkawiensis
C.astrum vs. C.lekaguli0.007< 0.001< 0.0010.0180.002
C.astrum vs. C.stellatus< 0.0010.0160.0490.009
C.langkawiensis vs. C.lekaguli0.014< 0.0010.042< 0.001
C.langkawiensis vs. C.stellatus0.0010.003< 0.0010.0010.002< 0.001
C.lekaguli vs. C.stellatus< 0.001< 0.0010.002< 0.0010.0420.030
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