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 333ED315-A455-5578-9D5E-3A3BC20D8AEB Cyrtodactylus uthaiensis https://zoobank.org/9422F241-3A46-4FAD-97C4-D6A6AF16B405 sp. nov.Fig. 12 Suggested Common Name: Uthai Thani Bent-toed GeckoHolotype.

Adult male ZMKU R 00949 from Thung Na Ngam Subdistrict, Lan Sak District, Uthai Thani Province, Thailand (15.37649°N, 99.63426°E, 106 m a.s.l.), collected by Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon, and Natee Ampai on 18 June 2018.

Diagnosis.

Cyrtodactylusuthaiensis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 13–15 supralabials, 10–11 infralabials, 33 paravertebral tubercles, 17 rows of longitudinally arranged tubercles, 36 transverse rows of ventrals, 159 longitudinal rows of ventrals, eight expanded subdigital lamellae on the fourth toe, 12 unexpanded subdigital lamellae on the fourth toe, 20 total subdigital lamellae on the fourth toe; seven expanded subdigital lamellae on the fourth finger, 11 unexpanded subdigital lamellae on the fourth finger, 18 total subdigital lamellae on the fourth finger; 16 total enlarged femoral scales, 12 total femoral pores; 14 enlarged pore-bearing precloacals; enlarged femorals and enlarged precloacals not continuous; proximal femorals less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and wide ventrolateral caudal fringe; ventrolateral caudal fringe composed scales of different size; tail circular in cross-section; slightly enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginated; maximum SVL 58.1 mm; and six dark transverse body bands (Tables 5, 6).

Description of holotype

(Fig. 12). Adult male SVL 58.1 mm; head moderate in length (HL/SVL 0.28), width (HW/HL 0.68), depth (HD/HL 0.39), distinct from neck, triangular in dorsal profile; lores concave slightly 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 0.29); ear opening elliptical, small; eye to ear distance greater than diameter of eye; rostral rectangular, dorsally furrowed, bordered posteriorly by large left and right supranasals, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by two smaller postnasals, ventrally by first supralabial; 13R/15L rectangular supralabials tapering smoothly to posterior margin of eye; 10R/11L infralabials tapering smoothly to posterior margin of eye; scales of rostrum and lores flat to domed, slightly larger than granular scales on top of head and occiput; scales of occiput intermixed with distinct, small tubercles; superciliaries subrectangular, largest dorsally and anteriorly; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting medially for approximately 40% of their length posterior to mental; one row of slightly enlarged, elongate sublabials extending posteriorly to fifth(L) and seventh(R) infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth, imbricate, pectoral and ventral scales.

Significant p-values from the results of the ANOVA analyses comparing all combinations of OTU pairs of the Cyrtodactylusbrevipalmatus group. SVL and TL4U are not listed because no species pairs differed significantly from one another. * = Results based on a Games-Howell post hoc test. Character abbreviations are listed in the Materials and methods.

Morphometric characters AG*HumL* ForL FemL TibL HL HW HD*ED*EE* ES EN* IO EL IN
Cyrtodactylusfluvicavus sp. nov. vs. C.brevipalmatus0.000.01< 0.0010.0010.048< 0.0010.0040.0020.004
C.interdigitalis vs. C.brevipalmatus0.000.000.00< 0.001
C.ngati vs. C.brevipalmatus0.000.0030.001< 0.001
C.ngati3 vs. C.brevipalmatus0.0140.030.01< 0.0010.0080.038< 0.0010.0040.000
C.rukhadeva vs. C.brevipalmatus< 0.0010.0010.022< 0.0010.021
C. sp.9 vs. C.brevipalmatus0.01< 0.001< 0.0010.03
C.interdigitalis vs. Cyrtodactylusfluvicavus sp. nov.0.01
C.ngati vs. Cyrtodactylusfluvicavus sp. nov.< 0.0010.0020.04< 0.0010.005< 0.001< 0.001
C.ngati3 vs. Cyrtodactylusfluvicavus sp. nov.< 0.0010.0250.038
C.rukhadeva vs. Cyrtodactylusfluvicavus sp. nov.0.049< 0.001< 0.0010.004
C. sp.9 vs. Cyrtodactylusfluvicavus sp. nov.0.00.0070.0130.0230.007
C.ngati vs. C.interdigitalis0.0070.0310.010< 0.0010.000
C.ngati3 vs. C.interdigitalis0.0110.003
C.rukhadeva vs. C.interdigitalis0.00.010.000.044
C. sp.9 vs. C.interdigitalis0.000.007
C.ngati3 vs. C.ngati< 0.0010.0190.006< 0.0010.001
C.rukhadeva vs. C.ngati0.003< 0.001< 0.0010.046< 0.0010.001< 0.001< 0.0010.001
C. sp.9 vs. C.ngati< 0.001< 0.0010.0420.007< 0.001< 0.0010.000
C.rukhadeva vs. C.ngati3< 0.0010.0210.00.000.0070.020.006
C. sp.9 vs. C.ngati30.0010.010.030.0030.0430.0010.0190.0190.003
C. sp.9 vs. C.rukhadeva0.020.0040.020.033
Meristic characters SL IL*PVT* LRT VS VSM TL4E TL4T FL4E FL4U*FL4T* FS PCS*BB*
Cyrtodactylusfluvicavus sp. nov. vs. C.brevipalmatus0.037< 0.0010.05
C.interdigitalis vs C.brevipalmatus< 0.0010.0000.0020.044
C.ngati vs C.brevipalmatus< 0.0010.028< 0.001
C.ngati3 vs C.brevipalmatus0.0170.029
C.rukhadeva vs C.brevipalmatus< 0.001< 0.001
C. sp.9 vs C.brevipalmatus< 0.0010.0030.022< 0.0010.05
C.interdigitalis vs Cyrtodactylusfluvicavus sp. nov.0.0000.0000.021< 0.0010.000< 0.001
C.ngati vs Cyrtodactylusfluvicavus sp. nov.0.0370.002< 0.001< 0.0010.0330.0000.002< 0.001
C.ngati3 vs Cyrtodactylusfluvicavus sp. nov.0.005
C.rukhadeva vs Cyrtodactylusfluvicavus sp. nov.0.0000.0010.001
C. sp.9 vs Cyrtodactylusfluvicavus sp. nov.< 0.001< 0.0010.0010.0040.020
C.ngati vs C.interdigitalis0.0010.029< 0.001< 0.0010.029
C.ngati3 vs C.interdigitalis0.0290.0250.005
C.rukhadeva vs C.interdigitalis0.0040.0110.0010.0440.0010.002
C. sp.9 vs C.interdigitalis0.003< 0.0010.0050.0430.01< 0.001
C.ngati3 vs C.ngati0.02670.001< 0.001
C.rukhadeva vs C.ngati< 0.001< 0.0010.008< 0.0010.0080.03< 0.001
C. sp.9 vs C.ngati0.0030.0160.0110.000< 0.001< 0.001< 0.001< 0.001
C.rukhadeva vs C.ngati3
C. sp.9 vs C.ngati30.0010.042< 0.0010.001
C. sp.9 vs C.rukhadeva0.029< 0.0010.0020.001
10.3897/zookeys.1129.90535.figure12ACF60A44-94B2-5F83-B4D0-2F637298548B

Holotype of Cyrtodactylusuthaiensis sp. nov. ZMKU R 00949 (field no. AA 06298) from Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam Subdistrict 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/769833

Body relatively short (AG/SVL 0.46) 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; smaller tubercles extend anteriorly onto nape and occiput, diminishing in size anteriorly; approximately 17 longitudinal rows of tubercles at midbody; approximately 33 paravertebral tubercles; small tubercles on flanks; 36 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 159 transverse rows of ventral scales; 16 total large femoral scales; 12 total femoral pores; 14 enlarged pore-bearing precloacals; no deep precloacal groove or depression; and two rows of post-precloacal scales on midline.

Forelimbs moderate in stature, relatively short (ForL/SVL 0.14); granular scales of forearm slightly larger than those on body, interspersed with large 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; 7R/7L expanded and 11R/11L 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; one row of 6R/6L of enlarged pore-bearing femoral scales not continuous with enlarged pore bearing precloacal scales, terminating distally at knee; 8R/8L enlarged femoral scales; 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; 8R/(broken)L wide, transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that extend onto sole, 12R/(broken)L unexpanded lamellae beneath first toe; and claws well-developed, sheathed by a dorsal and ventral scale at base.

Tail original, long 76.7 mm (TL/SVL 1.32), 4.1 mm in width at base, tapering to a point; sub-circular or nearly round 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 small but distinct ventrolateral caudal fringe; larger scales of ventrolateral fringe occur at regular intervals; slightly enlarged medial subcaudals intermittent, medially furrowed, posteriorly; single enlarged medial subcaudals absent; subcaudal scales, larger than dorsal caudal scales; base of tail bearing hemipenial swellings; 3R/3L conical postcloacal tubercles at base of hemipenial swellings; and postcloacal scales flat, imbricate.

Coloration in life

(Fig. 12). Ground color of the head, body, limbs, and tail pale-brown; dark, diffuse mottling on interorbital region and snout; dark blotch on top of head; wide, pale-colored postorbital stripe irregularly edged in dark brown extends from posterior margin of one eye across nape to posterior margin of other eye; ventral portion of lores, suborbital region, and supralabials darkly mottled; wide, dark brown nuchal band, bearing two posterior projections; six irregularly shaped darkly edged body bands extending between forelimb and hind limb insertions followed by one dark sacral band; paired dark brown paravertebral blotches on nape; band interspaces bearing irregularly shaped, dark-colored markings; dark-colored speckling on limbs and digits; digits bearing pale-colored bands; seven wide dark-colored caudal bands separated by six pale-colored bands; caudal bands encircle tail resulting in heavily mottled subcaudal region; all other ventral surfaces beige, generally immaculate; and iris gold in color.

Distribution.

Cyrtodactylusuthaiensis sp. nov. is currently known from the type locality at Thung Na Ngam Subdistrict, Lan Sak District, Uthai Thani Province, Thailand.

Etymology.

The specific epithet uthaiensis refers to the type locality, Uthai Thani Province, Thailand.

Comparisons.

Cyrtodactylusuthaiensis sp. nov. is the sister species to a clade comprised the sister species C.interdigitalis and C. sp.11 (Fig. 2). Together, these taxa form the sister lineage to C.cf.ngati1, C.cf.ngati2, and all other C.ngati. Cyrtodactylusuthaiensis sp. nov. differs from those lineages by an uncorrected pairwise sequence divergence of 5.81–8.13% (Table 2). We are aware that any comparison based on morphometric and meristic characters are preliminary being that there is only one sample of Cyrtodactylusuthaiensis sp. nov. and that additional sampling may preclude some characters and being diagnostic just as it may reveal that other characters are diagnostic (see Table 5). Therefore, at this point we rely on the invariable categorical characters to separate Cyrtodactylusuthaiensis sp. nov. from other species in the brevipalmatus group. Cyrtodactylusuthaiensis sp. nov. differs from C.brevipalmatus, Cyrtodactylusfluvicavus sp. nov., C.interdigitalis, C.ngati, C.ngati3, C.cf.ngati1, C.cf.ngati2, C.rukhadeva, C.cf.rukhadeva and C. sp.13 by having large dorsolateral caudal tubercles (DCT) forming a wide ventrolateral caudal fringe (VLF1). Cyrtodactylusuthaiensis sp. nov. is further differentiated from C.ngati3, C.cf.ngati1, C.cf.ngati2, C.interdigitalis, C.rukhadeva, C.cf.rukhadeva, and sp.13 by having a ventrolateral fringe not homogenous (VLF2). It differs from Cyrtodactyluskochangensis sp. nov., Cyrtodactylusrivularis sp. nov., C.rukhadeva, C.cf.rukhadeva, and C. sp.11 by having tail that is more circular in cross-section than square (TLcross). From Cyrtodactylusrivularis sp. nov., C.rukhadeva and C.cf.rukhadeva, it differs by having enlarged, unmodified, medial subcaudal scales (SC1). From Cyrtodactylusrivularis sp. nov., C.rukhadeva and C.cf.rukhadeva, it differs by lacking single, enlarged, medial subcaudal scales (SC2). From all species in the brevipalmatus group except C.interdigitalis it differs by having posteriorly emarginated, medial subcaudals bearing a median furrow (SC3).

Natural history.

Cyrtodactylusuthaiensis sp. nov. is the only species of the brevipalmatus group that occurs in an isolated hilly area within the Chao Phraya River Basin (Fig. 1). The holotype (ZMKU R 00949) was collected at night (2055 h) on a bamboo twig approximately 170 cm above ground level at 106 m elevation. The habitat was isolated karst formations within a mixed deciduous forest. This area was surrounded by agricultural fields (plantations and rice fields) and human residential areas (Fig. 13). The new species was found to co-occur with a gekkonid lizard, Dixoniussiamensis (Boulenger, 1899).

Summary statistics from the PERMANOVA analysis of the OTUs and proposed morphogroups of the Cyrtodactylusbrevipalmatus group.

OTU pairsF modelR2p-valuep-adjusted
C.rukhadeva vs. C.cf.ngati26.84740.40640.0150.544
C.rukhadeva vs. C.ngati38.88240.44670.0030.122
C.rukhadeva vs. C.interdigitalis3.36300.21890.0060.201
C.rukhadeva vs. C.ngati10.45800.48740.0030.114
C.rukhadeva vs. C.brevipalmatus6.59830.33670.0000.012
C.rukhadeva vs. Cyrtodactylusfluvicavus sp. nov.6.63570.30670.0000.004
C.rukhadeva vs. C. sp.93.86460.19450.0010.033
C.cf.ngati2 vs. C.brevipalmatus15.48180.75590.0481.000
C.cf.ngati2 vs. Cyrtodactylusfluvicavus sp. nov.15.91860.69460.0270.967
C.cf.ngati2 vs. C. sp.919.01300.70380.0220.804
C.ngati3 vs. C.interdigitalis4.47530.47230.0291.000
C.ngati3 vs. C.brevipalmatus14.94250.71350.0180.643
C.ngati3 vs. Cyrtodactylusfluvicavus sp. nov.8.79530.52370.0090.317
C.ngati3 vs. C. sp.914.79780.62180.0060.226
C.interdigitalis vs. C.ngati9.89760.66440.0291.000
C.interdigitalis vs. C.brevipalmatus4.56460.39470.0080.278
C.interdigitalis vs. Cyrtodactylusfluvicavus sp. nov.6.71200.42720.0030.124
C.interdigitalis vs. C. sp.95.65850.36140.0020.067
C.ngati vs. C.brevipalmatus7.48180.55500.0180.643
C.ngati vs. Cyrtodactylusfluvicavus sp. nov.22.82340.74050.0080.283
C.ngati vs. C. sp.917.01460.65400.0060.227
C.brevipalmatus vs. Cyrtodactylusfluvicavus sp. nov.17.85850.64100.0010.048
C.brevipalmatus vs. C. sp.99.39600.46070.0010.025
Cyrtodactylusfluvicavus sp. nov. vs. C. sp.98.20470.38690.0000.005
10.3897/zookeys.1129.90535.figure13545D896D-D647-5BD0-8BB7-A80F1C1B5F6C

Habitat of Cyrtodactylusuthaiensis sp. nov. at Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam Subdistrict.

https://binary.pensoft.net/fig/769834

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