Animalia Squamata Gekkonidae GrismerL. LeeAowpholAnchaleeYodthongSiripornAmpaiNateeTermprayoonKorkhwanAksornneamAkrachaiRujirawanAttapolIntegrative taxonomy delimits and diagnoses cryptic arboreal species of the Cyrtodactylusbrevipalmatus group (Squamata, Gekkonidae) with descriptions of four new species from ThailandZookeys15112022112910916210.3897/zookeys.1129.90535 7E67190E-06B0-564C-B4C8-E5A323705A73 Cyrtodactylus kochangensis https://zoobank.org/96DF655B-BBB7-4C06-9418-744EBCD14703 sp. nov.Fig. 10 Suggested Common Name: Ko Chang Bent-toed GeckoHolotype.

Adult female ZMKU R 00945 from Ko Chang Island, Ko Phayam Subdistrict, Mueang Ranong District, Ranong Province, Thailand (9.82411°N, 98.43999°E, 36 m a.s.l.), collected by Siriporn Yodthong, Natee Ampai, Attapol Rujirawan, and Piyawan Puanprapai on 8 July 2017.

Additional material examined.

Cyrtodactyluscf.kochangensis sp. nov. adult male THNHM 01667 from Khlong Naka Wildlife Sanctuary, Suk Samran District, Ranong Province, Thailand (~N 9.4596, E 98.5044, elevation unknown), collected by Yodchaiy Chuaynkern between 28 December 2000 and 2 January 2001.

Diagnosis

(based on the holotype).Cyrtodactyluskochangensis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 12 or 13 supralabials, nine infralabials, 34 paravertebral tubercles, 14 rows of longitudinally arranged tubercles, 35 transverse rows of ventrals, 172 longitudinal rows of ventrals, 8 or 9 expanded subdigital lamellae on the fourth toe, 11 or 12 unexpanded subdigital lamellae on the fourth toe, 19–21 total subdigital lamellae on the fourth toe; eight expanded subdigital lamellae on the fourth finger, ten unexpanded subdigital lamellae on the fourth finger, 18 total subdigital lamellae on the fourth finger; 12 total enlarged femoral scales; 12 enlarged precloacal scales; enlarged femoral and enlarged precloacal scales not continuous; proximal femoral scales less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and a wide ventrolateral caudal fringe not composed homogeneous scales; tail square in cross-section; slightly enlarged paired medial subcaudals not posteromedially furrowed; maximum SVL 60.1 mm; five dark transverse body bands (Tables 4, 5).

Description of holotype

(Fig. 10). Adult female SVL 60.1 mm; head moderate in length (HL/SVL 0.29), width (HW/HL 0.67), depth (HD/HL 0.38), distinct from neck, triangular in dorsal profile; lores flat anteriorly, weakly inflated posteriorly; prefrontal region slightly concave; canthus rostralis rounded; snout elongate (ES/HL 0.40), rounded in dorsal profile; eye large (ED/HL 0.24); ear opening subcircular, small; eye to ear distance greater than diameter of eye; rostral rectangular, furrowed dorsally, bordered posteriorly by large left and right supranasals and one slightly smaller azygous internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by two slightly smaller postnasals, bordered ventrally by first supralabial; 12R/13L rectangular supralabials tapering smoothly to below eye; 9R/9L infralabials tapering smoothly to below eye; scales of rostrum and lores domed, slightly larger than granular scales on top of head and occiput; scales of occiput intermixed with distinct, small tubercles; superciliaries subrectangular, largest anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapazoidal postmentals contacting medially for approximately 40% of their length posterior to mental; one row of six (R,L) slightly enlarged sublabials extending posteriorly fifth infralabials, subsequent sublabials much smaller; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth, imbricate, pectoral and ventral scales.

10.3897/zookeys.1129.90535.figure10CD090ADE-34AC-506C-A2AD-9D04153C1156

Holotype of Cyrtodactyluskochangensis sp. nov. ZMKU R 00945 (field no. AA 04439) from Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko Chang A dorsal view B ventral view C dorsal view of head D ventral view of femoral and precloacal regions E dorsal view of tail F ventral view of tail G holotype in life.

https://binary.pensoft.net/fig/769831

Body relatively short (AG/SVL 0.48) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with larger conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; tubercles of nape and occiput smaller than those on body; approximately 14 longitudinal rows of tubercles at midbody; approximately 34 paravertebral tubercles; tubercles on upper flanks smaller than those on dorsum; approximately 35 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; approximately 172 transverse rows of ventral scales; no pore-bearing, precloacal scales; 12 enlarged precloacal scales; no deep precloacal groove or depression; and approximately three rows of post-precloacal scales on midline.

Forelimbs moderate in stature, relatively short (ForL/SVL 0.13); granular scales of forearm slightly larger than those on body, interspersed with tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 8R/8L expanded and 10R/10L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.14), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, subimbricate, larger than dorsals; subtibial scales flat, imbricate; no pore-bearing femoral scales; 6R/6L enlarged femoral scales; enlarged femoral scales not contiguous with enlarged precloacal scales, terminating distally at knee; proximal femoral scales smaller than distal femorals, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; digits relatively long, well-developed, inflected at basal interphalangeal joints; 9R/8L wide, transversely expanded subdigital lamellae on the fourth toe proximal to joint inflection that extend onto sole, and 12R/11L unexpanded lamellae beneath the fourth toe; and claws well-developed, sheathed by a dorsal and ventral scale at base.

Tail original (but in two pieces), long 80.3 mm (TL/SVL 1.34), 4.0 mm in width at base, tapering to a point; nearly square in cross-section; dorsal scales flat, square bearing tubercles forming paravertebral rows and large tubercles forming a dorsolateral longitudinal row; slightly larger, posteriorly directed, semi-spinose tubercles forming wide distinct ventrolateral caudal fringe; scales of ventrolateral fringe generally interspersed at regular intervals with larger spinose scales; medial subcaudal scales paired, slightly enlarged; subcaudals, larger than dorsal caudal scales; base of tail bearing hemipenial swellings; one conical postcloacal tubercle at base of hemipenial swellings; and postcloacal scales flat, imbricate.

Coloration in life

(Fig. 10). Ground color of the head, body, limbs, and tail pale brown; faint, diffuse mottling on rostrum; lores darkly colored; wide, distinct, pale-colored postorbital stripe; nuchal band faint, bearing two posterior projections; three very faint, wide, irregularly shaped body bands between limb insertions edged in slightly darker brown; band interspaces bearing irregularly shaped, faint, dark-colored markings; dark-colored speckling on limbs and digits; digits bearing pale-colored bands; four wide, faint, dark-colored caudal bands separated by three pale-colored bands on original portion of tail; all caudal bands encircle tail; all ventral surfaces beige, generally immaculate; and iris orangish to coppery in color.

Variation.

The additional specimen (THNHM 01667) closely approximates the holotype in overall coloration and pattern except that it is more boldly marked. It has four dark-colored body bands as opposed to three and a complete original tail bearing eight dark-colored and seven pale-colored bands. The pale-colored postorbital stripe is slightly thinner and less distinct. Meristic and morphometric differences are listed in Table 5. Given its overall morphological and color pattern similarities and close geographic proximity to the holotype (~ 40 km), we consider this individual as C.cf.kochangensis sp. nov. pending genetic data.

Distribution.

Cyrtodactyluskochangensis sp. nov. is currently known only from the type locality at Ko Chang Island, Ko Phayam Subdistrict, Mueang Ranong District, Ranong Province, Thailand. The additional population of pending species status occurs in the Khlong Naka Wildlife Sanctuary, Suk Samran District Ranong Province.

Etymology.

The specific epithet kochangensis is in reference to the type locality, Ko Chang, Ranong Province, Thailand

Comparisons

(based on the holotype). Cyrtodactyluskochangensis sp. nov. forms a clade with the sister species Cyrtodactylusrivularis sp. nov. and C.rukhadeva (Fig. 2) from which it differs by an uncorrected pairwise sequence divergence of 12.00–12.52% and 12.52–13.68%, respectively (Table 2) and it and C.cf.kochangensis sp. nov. are separated from C.rukhadeva and Cyrtodactylusrivularis sp. nov. by geographic distance of no less than ~280–470 km (Fig. 1). The small sample size (N = 1) precludes it from statistical analyses, however at this point, it differs from C.rukhadeva and C.cf.rukhadeva in having 34 PVT versus 26–30; 14 LRT versus 18–20; 172 VSM versus 152–165; 12 FS versus 13–18; 12 PCS versus 13–17 and 1 PCT versus 2 or 3, collectively. From C.rukhadeva by having 12 FS versus 16 or 17; and five BB versus three. From Cyrtodactylusrivularis sp. nov. it differs in having 14 LRT versus 18–20; 172 VSM versus 160–166; 12 FS versus 14–16; and five BB versus three or four. Discrete differences between Cyrtodactyluskochangensis sp. nov. and C.cf.kochangensis sp. nov. and all other species and populations are presented in Tables 4, 5.

Natural history.

The holotype (ZMKU R 00945) was collected at night (2107 h) among branches of a small tree approximately 100 cm above the ground at 36 m elevation with a temperature of 28.6 °C and relative humidity of 83.9%. The surrounding habitat was dry evergreen forest with a rocky stream nearby (Fig. 11). The new species was found to co-occur with two other species of gekkonid lizards, Cyrtodactylusoldhami (Theobald, 1876) and Gekkotokehos (Grismer, Wood, Grismer, Quah, Thy, Phimmachak, Sivongxay, Seateun, Stuart, Siler, Mulcahy, Anamza & Brown, 2019).

10.3897/zookeys.1129.90535.figure1148F00AFB-8559-56D9-AEC0-41F79C090107

Habitat of Cyrtodactyluskochangensis sp. nov. at Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko Chang.

https://binary.pensoft.net/fig/769832

Summary statistics of the normalized morphometric data for the putative species of the Cyrtodactylusbrevipalmatus group. SD = ± standard deviation. Character abbreviations are listed in the Materials and methods.

Species SVL AG HumL ForL FemL TibL HL HW HD ED EE ES EN IO EL IN
C.brevipalmatus (N = 5)
Mean1.821.490.960.961.061.021.271.100.870.670.750.860.730.680.070.32
SD0.0200.0270.0310.0230.0240.0360.0120.0060.0170.0410.0250.0080.0190.0430.0580.041
Lower1.801.460.920.921.020.971.251.100.860.630.710.850.700.62-0.020.25
Upper1.851.520.990.981.091.071.291.110.890.730.770.870.750.730.130.36
C.elok (N = 4)
Mean1.911.590.801.051.141.131.341.201.000.710.830.950.790.700.190.43
SD0.0160.0120.3410.0260.0220.0240.0010.0120.0080.0170.0200.0080.0100.0700.0620.033
Lower1.891.570.341.021.111.101.341.180.980.680.810.940.780.610.150.40
Upper1.931.601.091.081.161.151.341.211.000.720.850.950.800.760.280.48
Cyrtodactylusfluvicavus sp. nov. (N = 7)
Mean1.861.520.951.011.111.031.311.140.910.690.780.920.790.730.200.38
SD0.0270.0090.0130.0170.0180.0120.0070.0110.0110.0070.0160.0080.0110.0060.040.02
Lower1.821.510.940.981.101.011.301.130.900.680.760.910.780.720.140.35
Upper1.891.540.981.041.151.051.321.160.930.700.810.940.810.740.240.40
C.interdigitalis (N = 4)
Mean1.861.500.981.011.101.081.301.120.800.720.760.900.770.680.130.33
SD0.0600.0160.0320.0150.0200.0070.0120.0060.1870.0260.0260.0240.0340.03070.0330.035
Lower1.781.480.940.991.071.071.281.120.560.690.730.880.740.660.100.29
Upper1.911.521.011.021.121.091.311.130.960.750.790.940.810.730.170.38
Cyrtodactyluskochangensis sp. nov. (N = 1)
Value1.781.511.000.951.051.021.301.140.910.690.760.910.780.680.050.36
C.cf.kochangensis sp. nov. (N = 1)
Value1.851.501.020.941.081.081.271.090.900.720.700.880.750.600.120.36
C.ngati (N = 3)
Mean1.831.470.910.991.061.051.311.080.850.570.760.860.810.74-0.120.43
SD0.0090.0020.0040.0070.0000.0060.0010.0030.0100.0350.0160.0150.0040.0090.0190.008
Lower1.821.470.910.981.061.041.311.080.840.550.740.850.800.73-0.130.42
Upper1.841.470.911.001.061.051.311.090.860.610.770.880.810.75-0.090.43
C.ngati3 (N = 3)
Mean1.881.580.941.001.121.071.321.120.950.690.810.930.800.770.080.41
SD0.0390.0010.0020.020.0010.0170.0070.0020.0140.0030.0040.0070.0060.0220.0160.01
Lower1.851.580.930.981.121.051.311.120.940.690.800.920.790.740.070.40
Upper1.921.580.941.021.121.091.331.120.960.690.810.940.800.780.100.42
C.ngati4 (N = 1)
Value1.871.500.871.021.131.071.331.110.910.720.790.940.830.750.510.43
C.cf.ngati1 (N = 1)
Value1.891.590.961.031.131.121.341.120.950.850.760.960.820.730.280.49
C.cf.ngati2 (N = 2)
Mean1.921.591.011.021.171.11.341.150.950.720.800.940.800.650.030.37
SD0.0350.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.000
Lower1.891.591.011.021.171.101.341.150.950.720.800.940.800.650.030.37
Upper1.941.591.011.021.171.101.341.150.950.720.800.940.800.650.030.37
Cyrtodactylusrivularis sp. nov. (N = 2)
Mean1.851.530.890.981.041.031.301.160.910.760.800.910.770.750.050.34
SD0.0250.010.0020.0070.0070.0050.0050.0010.0130.0070.0240.0050.0040.0250.0190.016
Lower1.831.530.890.971.031.031.291.160.910.750.790.910.770.740.040.33
Upper1.871.540.900.981.041.041.301.160.920.760.820.910.780.770.060.35
C.rukhadeva and C.cf.rukhadeva (N = 10)
Mean1.851.491.010.951.051.021.301.140.920.710.750.910.770.670.090.35
SD0.0260.0280.0550.0290.0270.0230.0090.0180.0250.0360.0340.0140.0150.0870.0690.022
Lower1.791.450.910.900.990.971.291.100.870.620.700.890.740.460.000.32
Upper1.881.541.081.001.091.051.321.160.950.750.800.920.790.730.230.38
Cyrtodactylusuthaiensis sp. nov. (N = 1)
Value1.761.800.950.991.091.021.281.100.760.720.740.880.750.680.190.32
C. sp.9 (Thong Pha Phum) (N = 8)
Mean1.861.530.910.971.081.021.301.160.890.710.770.900.770.740.090.34
SD0.0240.0160.0340.0210.0320.0190.0140.0170.0080.0150.0040.0120.0070.0120.0610.023
Lower1.811.500.860.931.020.991.271.140.880.690.760.890.760.720.000.30
Upper1.881.550.951.001.121.051.321.190.900.740.780.930.780.760.180.38
C. sp.11 (N = 1)
Value1.831.531.010.991.131.091.301.140.750.700.780.920.780.610.080.35
C. sp.13 (N = 2)
Mean1.831.450.940.941.071.011.271.110.910.670.750.880.760.740.150.34
SD0.0400.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.0000.000
Lower1.801.450.940.941.071.011.271.110.910.670.750.880.760.740.150.34
Upper1.861.450.940.941.071.011.271.110.910.670.750.880.760.740.150.34

Sex and raw meristic, categorical, and morphometric data used in the analyses from specimens in the Cyrtodactylusbrevipalmatus group. m = male; f = female; R/L = right/left; / = data unavailable.

Species C.brevipalmatus C.cf.brevipalmatus C.cf.brevipalmatus C.brevipalmatus C.brevipalmatus C.elok C.elok C.elok C.elok C.interdigitalis C.interdigitalis C.interdigitalis C.interdigitalis C. sp.11 C.ngati C.ngati C.ngati C.ngati C.ngati3C.ngati3C.ngati3C.ngati4C.cf.ngati1C.cf.ngati2C.cf.ngati2 C.rukhadeva C.rukhadeva C.rukhadeva C.cf.rukhadeva C.cf.rukhadeva C.cf.rukhadeva C.cf.rukhadeva C.cf.rukhadeva C.cf.rukhadeva C.cf.rukhadeva Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactylusfluvicavus sp. nov.Cyrtodactyluskochangensis sp. nov C.cf.kochangensis Cyrtodactylusuthaiensis sp. nov.Cyrtodactylusrivularis sp. nov.Cyrtodactylusrivularis sp. nov.C. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.9 Thong Pha PhumC. sp.13C. sp.13
Institutional catalog numberLSUHC 1899LSUHC 15076LSUHC 11788THNHM 10670THNHM 14112LSUHC 8238LSUHC 12180LSUHC 12181ZMMU R-16144THNHM 20226 paratypeTHNHM 20228 paratypeTHNHM 20229 paratypeTHNHM 20227 paratypeZMMU R-16492HNUE-R00111 holotypeIEBR 4829 paratypeVNUF R.2020.12 paratypeHNUE-R00112 paratypeFMNH 255454FMNH 270493FMNH 270492FMNH 265806NCSM 79472ZMMU R-14917NCSM 80100ZMMU R-16851 holotypeZMMU R-16852 paratypeZMKU R 00948 topotypeTHNHM 24622THNHM 24838THNHM 03251THNHM 03252THNHM 03253THNHM 03254THNHM 01807ZMKU R 00959 holotypeZMKU R 00958 paratypeZMKU R 00960 paratypeZMKU R 00961 paratypeZMKU R 00962 paratypeZMKU R 00963 paratypeZMKU R 00964 paratypeZMKU R 00945 holotypeTHNHM 01667ZMKU R 00949 holotypeZMKU R 00947 holotypeZMKU R 00946 paratypeZMKU R 00950ZMKU R 00951ZMKU R 00952ZMKU R 00953ZMKU R 00954ZMKU R 00955ZMKU R 00956ZMKU R 00957THNHM 00104THNHM 27821
Sexmfffffmmfffffmmffffmmmfffmffmfmmfmmmmmfffffmmfffmfmmfmfff
Meristic data
Supralabials (SL)11121014121181391412111211101010101313131014912119141113131112131212R/12L13R/12L13R/12L11R/12L12R/12L13R/12L12R/11L12R/13L1213R/15L13R/12L13R/12L12131314131313131215
Infralabials (IL)81091111118119988799999109118111012101191010101010111010R/10L10R/10L9R/10L10R/10L10R/10L10R/10L10R/10L9R/9L1010R/11L11R/10L10R/9L8810109101091011
Paravertebral tubercles (PVT)39373837370000323333333039403840282726272832292730302628272730302630282727282628342933343332333434363630303329
Longitudinal rows of tubercles (LRT)15161716146744192019191818181722191817191824191920191819181819191917171416171816141917201821192020212119191820
Ventral scales (VS)383838363945454736424042433438363532373636333336353443383836373739343534373330363739353436343734333334303332333736
Ventral scales along middle of the body (VSM)176170182154160190225234192187170187178160168164178158159166156158164166165154152165162158157159168160161155154155172164175170172159159160166173158156166159159150169159165
Expanded subdigital lamellae on 4th toe (TL4E)7898810999121010119810991010810981099989910910109R/9L10R/10L9R/9L9R/9L10R/11L9R/10L9R/9L9R/8L88R/(broken)L9R/9L9R/9L910981089997
Unmodified subdigital lamellae on 4th toe (TL4U)13111111121110119141312141011101110111111111210101111121113121215131311R/11L12R/11L10R/10L12R/12L11R/11L10R/10L12R/13L12R/11L1312R/(broken)L13R/13L12R/13L12141312131211131212
Total subdigital lamellae 4th toe (TL4T)201920192021192018262322231913161716212119212118202018211922212214232320R/20L22R/21L19R/19L21R/21L21R/22L19R/20L22R/22L21R/19L212022R/22L21R/22L21242220232020222119
Expanded subdigital lamellae on 4th finger (FL4E)8887889999899106676888897998878888888R/8L8R/8L8R/8L8R/8L7R/7L8R/9L7R/7L8R/8L87R/7L8R/8L8R/8L8778888888
Unmodified subdigital lamellae on 4th finger (FL4U)9111010101213981211121399999101010108910109111011101012121210R/10L10R/10L10R/9L11R/11L10R/10L9R/9L10R/10L10R/10L1211R/11L11R/10L12R/12L10121211121211121110
Total subdigital lamellae 4th finger (FL4T)171918171820221817212121221915151815181818181716191917191717181820202018R/18L18R/18L18R/17L19R/19L17R/17L17R/18L17R/17L18R/18L2018R/18L19R/18L20R/20L18191919202019201918
Enlarged femoral scales (R/L)0008R/8L7R/7L000011R/8L10R/9L8R/8L9R/10L9R/8L10R/10L9R/8L10R/9L8R/9L9R/7L8R/9L9R/9L8R/8L9R/8L7R/8L7R/8L9R/8L8R/8L9R/8L9R/L9R/9L9R/7L7R/7L6R/7L5R/8L7R/7L5R/6L4R/5L5R/6L6R/6L5R/6L5R/6L6R/6L6R/6L7R/7L8R/8L8R/8L6R/8L5R/7L8R/8L8R/8L7R/8L8R/8L7R/8L7R/6L8R/8L9R/9L7R/10L
Total enlarged femoral scales (FS)1610111614000014191919172017191716171816171515171617181816141313141191112111112121416161412161615161513161817
Total femoral pores in males (FP)70000000000001714000014151300017001401213011131181000000141200016014151412000
Enlarged precloacal scales (PCS)777878887141513191313131313151313131213131713151515141315151415141415141515121614151517151515151515151416
Precloacal pores in males (PP)7000008800000130000131313130001713015014130151415141400000161400015015151515000
Postcloacal tubercles (PCT)3323332333233332120000234322R/3L32322323R/2L3R/2L3R/3L1R/1L3R/2L3R/3L2R/2L1R/1L33R/3L2R/2L3R/3L22R/3L332R/3L2R/3L3233
Body bands (BB)46355553355553666634333333333344//5333333355634343435443/
Categorical data
Small tubercles on flank (FKT)presentpresentpresentpresentpresentabsentabsentabsentabsentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresentpresent
Dorsolateral caudal tubercles (DCT)smallsmallsmall/smalllargelargelargelargesmall/smallsmalllargesmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmallsmall/smallsmallsmallsmallsmallsmallsmalllargelargelargelargelargelargelargelargelargelargelarge/largesmallsmall
Ventrolateral caudal fringe narrow or wide (VLF1)narrownarrownarrow/narrowwidewidewidewidenarrow/narrownarrowwidenarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrownarrow/narrownarrownarrownarrownarrownarrownarrowwidewidewidewidewidewidewidewidewidewidewide/widenarrownarrow
Ventrolateral caudal fringe scales generally homogenous (VLF2)nonono/nononononoyesyesyesyesyesnonononoyesyesyesyesyesyesyesyesyesyesyesyesyesyesyesyes/nonononononononononoyesyesnononononono/noyesyes
Tail cross-section (TLcross)circularcircularcircular/circularsquaresquaresquaresquarecircular/circularcircularsquarecircularcircularcircularcircularcircularcircularcircularcircularcircularcircularcircularsquaresquaresquaresquaresquaresquaresquaresquaresquare/circularcircularcircularcircularcircularcircularcircularsquare/circularsquaresquaresquaresquaresquaresquaresquaresquare/squarecircularcircular
Slightly enlarged medial subcaudals (SC1)presentpresentpresent/absentabsentabsentabsentabsentabsent/absentabsentpresentpresentpresentpresentpresent/presentpresentpresentpresentpresentpresentabsentabsentabsentabsentabsentabsentabsentabsentabsent/presentpresentpresentpresentpresentpresentpresentpresentpresentpresentabsentabsentpresentpresentpresentpresentpresentpresent/presentpresentpresent
Single enlarged medial subcaudal (SC2)absentabsentabsent/absentabsentabsentabsentabsentabsent/absentabsentabsentabsentabsentabsentabsent/absentabsentabsentabsentabsentabsentpresentpresentpresentpresentpresentpresentpresentpresentpresent/absentabsentabsentabsentabsentabsentabsentabsentabsentabsentpresentpresentabsentabsentabsentabsentabsentabsent/absentabsentabsent
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3)nonono/nononononoyes/yesyesnonononono/nononononononononononononononononononononononononoyesnonononononoyesno/nonono
Morphometric data
SVL 68.870.864.165.9563.7980.278.284.878.681.1974.8078.5659.7068.166.568.169.346.683.670.274.173.878.087.177.774.971.771.668.371.873.675.374.773.261.572.572.069.668.476.865.778.260.170.258.173.968.173.173.573.773.264.476.676.674.263.772.9
AG 35.733.430.130.026.539.737.841.536.234.533.732.724.634.628.829.830.219.741.335.437.031.338.241.936.834.632.633.927.329.930.931.332.230.326.233.433.632.030.435.630.638.129.031.526.634.833.234.833.935.433.628.537.133.235.125.830.6
HumL 9.79.38.09.69.510.29.110.11.79.810.211.27.410.37.98.18.55.68.68.78.66.98.711.59.210.710.47.99.88.312.211.311.811.010.19.18.89.08.010.07.510.16.510.27.08.17.68.47.29.09.07.28.08.18.67.610.1
ForL 9.99.88.98.28.711.511.711.810.210.610.511.18.48.59.210.010.16.510.29.310.410.010.310.410.78.67.99.68.78.59.010.69.69.27.910.510.310.510.111.18.810.87.68.68.39.79.19.59.19.29.89.210.08.69.88.19.6
FemL 12.012.611.511.79.812.914.214.613.114.713.212.710.212.611.511.511.57.613.712.713.013.113.115.214.212.611.810.510.810.911.510.211.912.19.513.112.512.513.514.111.513.910.412.110.011.410.412.811.612.312.510.913.710.812.510.712.8
TibL 11.612.210.59.78.213.514.013.812.313.111.912.910.211.410.811.111.87.812.511.811.211.112.812.612.710.19.311.29.710.710.911.711.311.19.111.310.610.29.911.29.412.38.411.88.411.210.310.510.110.610.69.911.110.011.410.110.2
HL 19.319.319.017.918.221.821.621.921.720.819.921.716.718.420.120.420.716.121.720.620.320.721.222.121.420.219.219.719.719.920.821.320.821.517.920.120.519.720.121.218.621.317.318.316.120.319.319.920.920.120.017.620.419.320.017.619.9
HW 13.213.812.312.312.015.616.115.915.114.013.414.211.413.112.612.011.88.813.812.513.012.312.714.813.514.613.414.013.113.914.915.013.114.111.814.013.412.913.014.913.015.411.612.110.914.913.714.514.315.713.912.814.714.414.111.913.8
HD 8.07.67.67.37.09.69.810.49.83.48.68.76.68.37.47.26.65.19.28.49.17.68.38.79.29.28.58.37.38.98.28.28.18.97.58.58.18.37.98.17.88.36.57.86.38.28.27.87.77.97.77.08.27.87.67.78.4
ED 5.24.54.35.34.44.85.05.75.05.35.55.94.44.43.84.13.42.64.94.94.94.86.54.66.04.64.35.54.95.15.85.45.05.54.75.05.04.94.75.14.55.34.25.24.65.85.65.05.15.05.04.85.65.34.94.15.3
EE 5.75.94.95.75.76.47.17.06.85.86.26.44.86.25.85.55.94.46.96.16.25.75.36.56.26.26.25.85.16.25.65.75.46.24.36.55.95.75.86.15.46.55.04.94.76.56.25.95.96.05.95.36.16.06.04.96.3
ES 7.47.67.07.07.28.68.79.58.68.37.89.16.87.77.57.66.95.09.08.38.38.28.78.88.48.37.77.97.48.18.48.88.18.67.38.58.38.28.19.27.39.36.97.56.48.37.97.98.57.97.97.38.27.97.97.28.0
EN 5.75.44.95.35.46.06.26.56.26.05.56.85.15.56.76.36.24.56.56.26.16.26.26.66.06.35.75.85.46.06.26.45.86.25.36.56.25.96.16.65.66.55.25.54.96.15.86.06.16.05.85.46.16.05.95.65.9
IO 5.44.74.74.25.25.75.45.43.94.84.75.54.32.95.65.45.64.26.65.65.45.14.93.55.73.33.15.64.54.75.65.75.75.64.25.55.45.35.15.65.05.64.24.04.35.85.55.45.55.85.54.95.75.65.34.86.1
EL 1.01.41.11.31.01.91.41.51.41.31.31.61.20.90.80.80.70.31.31.11.21.01.51.20.91.21.01.41.61.51.21.31.21.20.91.41.51.71.41.81.61.81.01.31.51.11.11.11.51.51.21.21.01.21.31.41.4
IN 1.72.12.32.12.22.72.62.53.12.12.22.51.82.32.82.62.62.02.82.52.52.32.72.22.52.22.12.12.02.22.42.52.42.32.02.32.42.52.32.32.32.61.92.21.82.32.02.32.42.22.02.02.32.22.22.12.3
10.3897/zookeys.1129.90535.figure2CECE1A10-696A-5A3D-9B7A-FF98A8E3B584

Maximum clade credibility BEAST phylogeny of the Cyrtodactylusbrevipalmatus group highlighting the new species described herein. Bayesian posterior probabilities (BPP) are listed at the nodes.

https://binary.pensoft.net/fig/769823

Mean (minimum–maximum) percentages of uncorrected pairwise sequence divergence (p-distances) among the putative species of the Cyrtodactylusbrevipalmatus group based on 1,386 base pairs of mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs. Intraspecific p-distance are in bold font. n/a = data not applicable.

C.brevipalmatus C.cf.ngati1C.cf.ngati2 C.elok Cyrtodactylusfluvicavus sp. nov. C.interdigitalis Cyrtodactyluskochangensis sp. nov.C.ngati, C.ngati3 and C.ngati4Cyrtodactylusrivularis sp. nov. C.rukhadeva C. sp.9 (Thong Pha Phum)C. sp.10C. sp.11C. sp.14Cyrtodactylusuthaiensis sp. nov.
C.brevipalmatus N = 1 n/a
C.cf.ngati1 N = 121.03 n/a
C.cf.ngati2 N = 121.684.39 n/a
C.elok N = 120.7722.5821.42 n/a
Cyrtodactylusfluvicavus sp. nov. N = 718.8610.6411.0220.15 0.10
(18.84–18.97)(10.58–10.84)(10.97–11.23)(20.13–20.26) (0.00–0.26)
C.interdigitalis N = 120.776.979.1622.8412.02 n/a
(12.00–12.13)
Cyrtodactyluskochangensis sp. nov. N = 119.3514.5814.7120.9012.3115.23 n/a
(12.26–12.31)
C.ngati, C.ngati3 and C.ngati4 N = 720.703.303.7121.1111.348.1314.58 0.84
(20.65–20.90)(2.84–4.00)(3.35–4.26)(20.90–21.42)(11.10–11.87)(7.74–8.65)(14.45–14.84) (0.00–1.55)
Cyrtodactylusrivularis sp. nov. N = 220.0015.8715.0321.6112.5715.4812.2615.03 0.52
(19.74–20.26)(15.61–16.13)(14.84–15.23)(21.42–21.81)(12.26–13.03)(15.23–15.74)(12.00–12.52)(14.71–15.48)
C.rukhadeva N = 220.6515.4215.4821.6112.2516.0013.1015.234.65 1.55
(20.13–21.16)(14.84–16.00)(14.84–16.13)(21.16–22.06)(11.61–13.03)(15.35–16.65)(12.52–13.68)(14.19–16.23)(3.61–5.68)
C. sp.9 (Thong Pha Phum) N = 920.347.939.5122.029.758.9613.228.8113.1213.25 0.22
(20.13–20.65)(7.74–8.00)(9.42–9.55)(21.81–22.32)(9.55–9.94)(8.77–9.03)(13.03–13.29)(8.13–9.68)(12.77–13.42)(12.52–13.94) (0.00–0.52)
C. sp.10 N = 119.879.2910.8421.9410.1210.1913.6810.2113.9414.328.06 n/a
(10.06–10.32)(10.06–10.45)(13.68–14.19)(13.68–14.97)(7.87–8.13)
C. sp.11 N = 120.397.238.9022.1911.123.8714.588.2815.35 1)15.618.9610.45 n/a
(11.10–11.23)(8.00–8.65)(15.10–15.6(14.97–16.26)(8.77–9.03)
C. sp.14 N = 16.4520.9020.6520.0018.3420.1319.1020.5219.7420.0019.6018.8419.61 n/a
(18.32–18.45)(20.26–20.65)(19.48–20.00)(19.48–20.52)(19.48–19.87)
Cyrtodactylusuthaiensis sp. nov. N = 119.745.818.1321.1610.127.113.946.9713.9413.947.808.396.5819.48 n/a
(10.06–10.32)(6.58–7.61)(13.68–14.19)(13.29–14.58)(7.61–7.87)
10.3897/zookeys.1129.90535.figure13F3531E5-1850-5288-B05B-E07E00B8515B

Distribution of nominal species and unnamed populations and specimens of the Cyrtodactylusbrevipalmatus group. Stars denote type localities. White circles are literature localities from which specimens were not examined and remain unidentified. Locality data for all material examined is in Table 1.

https://binary.pensoft.net/fig/769822

Cyrtodactylus specimens examined in this study. Institutional abbreviations follow Sabaj (2020) except that YC = Yodchaiy Chuaynkern from Department of Biology, Faculty of Science, Khon Kaen University, Khon Kaen, Thailand.

SpeciesLocationCatalog no.GenBank no.
C.brevipalmatus Thailand, no dataLSUHC 1899not in tree
C.brevipalmatus Thailand, Nakhon Si Thammarat Province, Nopphitam District, Khao Nan National Park, Huay Lak Protected UnitTHNHM 10670not in tree
C.brevipalmatus Thailand, Nakhon Si Thammarat Province, Lan Saka District, Khao Luang National ParkTHNHM 14112not in tree
C.brevipalmatus Thailand, Nakhon Si Thammarat Province, Khao Ram Mt.AUP-00573 OK626313
C.cf.brevipalmatus Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung RayaLSUHC 11788not in tree
C.cf.brevipalmatus Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung RayaLSUHC 15076not in tree
C.elok Peninsular Malaysia, Pahang State, Fraser’s Hill, the GapZRC 2.6091/LSUHC 6471 JQ889180
C.elok Peninsular Malaysia, Negeri Sembilan StateLSUHC 8238not in tree
C.elok Peninsular Malaysia, Pahang State, near Cameron HighlandsLSUHC 12180not in tree
C.elok Peninsular Malaysia, Pahang State, near Cameron HighlandsLSUHC 12181not in tree
C.elok Malaysian pet trade, no dataZMMU R-16144not in tree
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00958 paratype OP620036
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00959 holotype OP620037
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00960 paratype OP620038
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00961 paratype OP620039
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00962 paratype OP620040
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00963 paratype OP620041
Cyrtodactylusfluvicavus sp. nov.Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National ParkZMKU R 00964 paratype OP620042
C.interdigitalis Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam NaoTHNHM 20226 paratypenot in tree
C.interdigitalis Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam NaoTHNHM 20227 paratypenot in tree
C.interdigitalis Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam NaoTHNHM 20228 paratypenot in tree
C.interdigitalis Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam NaoTHNHM 20229 paratypenot in tree
C.interdigitalis Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam NaoYC000952 ON055281
Cyrtodactyluskochangensis sp. nov.Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko ChangZMKU R 00945 holotype OP620023
C.cf.kochangensis sp. nov.Thailand, Ranong Province, Khlong Nakha Wildlife SanctuaryTHNHM 01667not in tree
C.ngati Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom CaveHNUE-R00111 holotype ON411220
C.ngati Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom CaveHNUE-R00112 paratypenot in tree
C.ngati Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom CaveIEBR 4829 paratype OK626318
C.ngati Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom CaveVNUF R.2020.12 paratype OK626319
C.ngati3Laos, Khammouane ProvinceVNUF R.2014.50 ON411221
C.ngati3Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation AreaFMNH 255454 JQ889181
C.ngati3Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation AreaFMNH 270492 OK626315
C.ngati3Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation AreaFMNH 270493not in tree
C.ngati4Thailand, Loei Province, Nam San Noi River, Phu Luang Wildlife SanctuaryFMNH 265806 JX519471
C.cf.ngati1Laos, Xaignabouli Province, Ban Pha Liep, Houay Liep StreamNCSM 79472 OK626316
C.cf.ngati2Laos, Vientiane ProvinceZMMU R-14917not in tree
C.cf.ngati2Laos, Vientiane Province, tributary of Nam Pha River, Houay Wan StreamNCSM 80100 OK626317
Cyrtodactylusrivularis sp. nov.Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U WaterfallZMKU R 00946 paratype OP620024
Cyrtodactylusrivularis sp. nov.Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U WaterfallZMKU R 00947 holotype OP620025
C.rukhadeva Thailand, Ratchaburi Province, Suan Phueng District, Khao Laem MountainZMMU R-16851 holotype OK626320
C.rukhadeva Thailand, Ratchaburi Province, Suan Phueng District, Hoop Phai TongZMMU R-16852 paratypenot in tree
C.rukhadeva Thailand, Ratchaburi Province, Suan Phueng District, Suan Phueng SubdistrictZMKU R 00948 OP620026
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 01807not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 03251not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 03252not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 03253not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 03254not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 24622not in tree
C.cf.rukhadeva Thailand, Phetchaburi Province, Kaeng Krachan National ParkTHNHM 24838not in tree
Cyrtodactylusuthaiensis sp. nov.Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam SubdistrictZMKU R 00949 holotype OP620027
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Thong Pha Phum National ParkAUP-01715 MT468909
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00950 OP620028
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00951 OP620029
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00952 OP620030
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00953 OP620031
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00954 OP620032
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00955 OP620033
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00956 OP620034
C. sp.9Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National ParkZMKU R 00957 OP620035
C. sp.10Thailand, Tak Province, Tha Song Yang District, Mae Moei National Park, Chao Doi WaterfallAUP-00680 MT468902
C. sp.11 (C.cf.interdigitalis)Thailand, Phitsanulok Province, Phu Hin Rong Kla National ParkZMMU R-16492 MW792061
C. sp.13Thailand, Tak Province, Umphang District, Thung Yai Naresuan Wildlife SanctuaryTHNHM 00104not in tree
C. sp.13Thailand, Kanchanaburi Province, Thong Pha Phum District, Ban Saphan LaoTHNHM 27821not in tree
C. sp.14 (C.cf.brevipalmatus)Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung RayaUSMHC 2555 OK626314