Corresponding author: Anchalee Aowphol (
Academic editor: Thomas Ziegler
The bent-toed geckos of the
Termprayoon K, Rujirawan A, Ampai N, Wood Jr PL, Aowphol A (2021) A new insular species of the
Bent-toed geckos in the genus
One clade of particular interest is the
Field surveys were conducted on Tarutao Island, Mueang Satun District, Satun Province, southern Thailand from November 2017 to November 2019 (Fig.
Map showing the type locality of
Total genomic DNA was extracted from ethanol-preserved liver tissue of five
Samples used in the molecular analyses, including their GenBank accession number (
Species | Locality | Museum No. | GenBank Accession No. | Reference |
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Pakistan, Baluchistan Province, Makran District, Gwadar division | FMNH 247474 |
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Unknow | NC 00155 |
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captive | JB 28 |
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Vietnam, Kien Giang Province, Kien Hai District, Hon Tre Island |
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Lao, Khammouan Province, Nakai District | FMNH 255454 |
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FMNH 265806 |
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BYU 53435 (holotype) |
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BYU 53436 (paratype) |
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BYU 53437 (paratype) |
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Phylogenetic trees were reconstructed using two different methods, Maximum Likelihood (
The morphological characters and their definition used in this study were modified from previous studies of the
Measurement and meristic characters used in this study, with abbreviations and explanations.
Abbreviations | Characters |
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Snout–vent length, taken from the tip of snout to the vent |
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Tail width, taken at the base of the tail immediately posterior to the postcloacal swelling |
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Tail length, taken from vent to the tip of the tail, original or regenerated |
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Forearm length, taken from the posterior margin of the elbow while flexed 90º to the inflection of the flexed wrist |
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Tibia length, taken from the posterior surface of the knee while flexed 90º to the base of the heel |
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Axilla to groin length, taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body |
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Head length, the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout |
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Head width, measured at the angle of the jaws |
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Head depth, the maximum height of head from the occiput to the throat |
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Eye diameter, the greatest horizontal diameter of the eyeball |
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Eye to ear distance, measured from the anterior edge of the ear opening to the posterior edge of the eyeball |
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Eye to snout distance, measured from anterior most margin of the eyeball to the tip of snout |
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Eye to nostril distance, measured from the anterior margin of the eyeball to the posterior margin of the external nares |
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Inter orbital distance, measured between the anterior edges of the orbit |
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Ear length, the greatest vertical distance of the ear opening |
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Internarial distance, measured between the nares across the rostrum |
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Supralabial scales, counted from the largest scale immediately posterior to the dorsal inflection of the posterior portion of the upper jaw to the rostral scale |
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Infralabial scales, counted from the largest scale immediately posterior to the dorsal inflection of the posterior portion of the upper jaw to the mental scale |
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The number of paravertebral tubercles between limb insertions, counted in a straight line immediately left or right of the vertebral column |
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The number of longitudinal rows of body tubercles, counted transversely across the center of the dorsum from one ventrolateral fold to the other |
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The number of longitudinal rows of ventral scales, counted transversely across the center of the abdomen from one ventrolateral fold to the other |
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The number of subdigital lamellae beneath the fourth toe, counted from the base of the first phalanx to the claw |
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The total number of precloacal and femoral pores in male (i.e., the sum of the number of femoral and precloacal scales bearing pores combined as a single meristic referred to as the femoroprecloacal pores) |
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The number of precloacal pores in female |
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The number of dark body bands between limb insertions |
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The number of dark caudal bands on the original tail |
External morphological characters examined in the
All analyses were performed using the base statistical software in R v3.6.1 (
Principal components analysis (
The aligned matrix contained 1,429 mtDNA characters from 93 individuals of the
The topologies of
Percentage uncorrected pairwise genetic divergence (
Species |
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1 | 2 | 3 | 4 | 5 | |
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1 | 5 |
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2 |
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12 | 10.50 (9.78–12.37) |
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3 |
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3 | 9.90 (9.56–10.88) | 9.86 (9.51–11.21) |
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4 |
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10 | 10.49 (9.86–11.69) | 10.19 (9.71–11.59) | 7.62 (7.39–7.83) |
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5 |
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9 | 9.33 (8.46–10.80) | 9.94 (8.98–11.77) | 8.58 (8.00–9.59) | 9.39 (8.42–10.54) |
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The Maximum Likelihood tree of the
The Bayesian consensus tree of the
The
Summary statistics and factor loadings of the first three principal components (
Characters | Males | Females | ||||
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PC2 | PC3 |
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PC2 | PC3 | |
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0.660 | -0.284 | 0.293 | 0.374 | -0.097 | -0.127 |
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0.436 |
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-0.073 | 0.380 |
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0.194 |
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-0.228 | -0.051 | 0.693 | 0.111 | 0.003 |
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0.098 | -0.032 |
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0.181 | 0.102 |
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-0.261 | - |
0.364 | -0.030 | 0.157 | 0.824 |
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0.459 | -0.508 | -0.560 | 0.532 | -0.194 | 0.537 |
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-0.168 | 0.177 |
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-0.350 | -0.197 |
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-0.325 | 0.188 | 0.526 | -0.647 | 0.170 |
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0.552 | -0.195 | -0.491 | 0.137 | -0.086 | 0.612 |
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-0.171 | 0.194 |
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-0.410 | 0.266 |
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0.113 | 0.054 |
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-0.078 | -0.271 |
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0.246 | 0.142 |
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0.122 | -0.263 |
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0.143 |
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0.435 | 0.361 |
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0.033 |
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0.303 |
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-0.417 | 0.343 | 0.690 | -0.298 |
INadj | -0.105 | 0.093 | 0.560 | -0.089 |
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0.238 |
Eigenvalue | 6.882 | 3.161 | 1.587 | 5.222 | 3.289 | 1.835 |
Percentage of variance | 45.879 | 21.073 | 10.581 | 34.813 | 21.928 | 12.235 |
Cumulative proportion | 45.879 | 66.952 | 77.533 | 34.813 | 56.741 | 68.976 |
Plots of the first two principal components of
The univariate analyses (
Summary pairwise results of statistically significant characters (Tukey’s test;
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M | F | M | F | M | F | ||
M | – | – | – | – | – | – | |
F | – | – | – | – | – | – | |
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M | – | – | – | – | – | |
F | – | – | – | – | – | ||
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M | – | – | – | – | – | – |
F | – | – | – | – | |||
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M | – | – | – | – | ||
F | – | – | – |
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Adult male (
Adult male holotype of
Head of the holotype of
Male holotype of
Male paratypes of
Female paratypes of
Adult male
Body relatively short (
Forelimbs moderate in stature, relatively short (
Precloacal region of
Tail 94.8 mm in length, completely regenerated, 9.2 mm in width at base, tapering to a point; regenerated tail covered with small, smooth, rectangular scales dorsally; base of tail bearing hemipenial swellings; one row of 4/4 medium-sized postcloacal tubercles on each hemipenial swelling; postcloacal scales smooth, flat, large, imbricate.
Descriptive measurement (millimeters), meristic (left/right) and color pattern characters of the type series of
H | P | P | P | P | P | P | P | P | P | |
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Sex | M | M | M | M | M | F | F | F | F | F |
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94.2 | 95.9 | 94.6 | 87.9 | 86.3 | 96.1 | 93.6 | 94.8 | 90.4 | 86.6 |
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94.8r | 96.8r | 92.3r | 69.0r | 18.5b | 124.4 | 70.5r | 107.5r | 81.7r | 102.3r |
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9.2 | 9.7 | 9.2 | 8.4 | 7.5 | 7.6 | 9.1 | 9.1 | 10.3 | 7.1 |
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15.4 | 15.3 | 15.1 | 14.4 | 14.1 | 14.9 | 14.5 | 15.9 | 15.6 | 14.8 |
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18.1 | 18.3 | 17.8 | 17.4 | 17.6 | 17.7 | 17.3 | 18.1 | 17.8 | 16.5 |
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43.3 | 43.7 | 43.9 | 44.4 | 39.6 | 46.0 | 44.5 | 44.5 | 45.4 | 39.9 |
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27.6 | 26.3 | 27.0 | 25.2 | 24.8 | 26.5 | 26.9 | 27.3 | 26.4 | 25.7 |
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16.7 | 16.3 | 17.3 | 15.4 | 14.7 | 16.4 | 16.8 | 17.2 | 15.7 | 15.2 |
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10.4 | 9.6 | 10.8 | 9.2 | 8.7 | 10.4 | 10.2 | 10.1 | 9.4 | 9.2 |
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6.8 | 6.7 | 6.7 | 5.9 | 5.8 | 6.4 | 6.8 | 6.6 | 6.6 | 5.6 |
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7.1 | 7.6 | 7.2 | 6.3 | 6.2 | 6.9 | 7.0 | 7.0 | 6.7 | 6.8 |
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10.8 | 10.7 | 10.9 | 9.8 | 9.8 | 10.9 | 10.8 | 10.7 | 10.4 | 10.1 |
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8.5 | 8.3 | 8.3 | 7.8 | 7.5 | 8.2 | 8.3 | 8.5 | 8.1 | 7.9 |
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5.9 | 6.2 | 6.6 | 5.8 | 5.5 | 6.0 | 6.4 | 6.1 | 5.7 | 6.2 |
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2.4 | 2.5 | 2.2 | 2.0 | 2.0 | 2.0 | 2.8 | 2.7 | 2.1 | 2.3 |
IN | 2.9 | 3.2 | 3.5 | 3.1 | 3.0 | 3.8 | 3.3 | 3.2 | 3.2 | 2.6 |
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13/14 | 14/13 | 15/12 | 13/14 | 13/12 | 13/13 | 13/13 | 13/12 | 12/12 | 13/13 |
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11/11 | 10/10 | 11/12 | 12/12 | 11/11 | 11/11 | 12/11 | 12/12 | 10/11 | 11/11 |
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36 | 35 | 41 | 38 | 43 | 38 | 40 | 40 | 40 | 47 |
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21 | 20 | 21 | 19 | 22 | 20 | 21 | 19 | 22 | 22 |
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33 | 37 | 36 | 35 | 36 | 37 | 39 | 37 | 37 | 36 |
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21/22 | 21/21 | 21/21 | 21/20 | 23/22 | 22/22 | 21/21 | 22/23 | 22/20 | 20/20 |
27 | 25 | 24 | 29 | 27 | / | / | / | / | / | |
/ | / | / | / | / | 15 | 12 | 14 | 11 | 11 | |
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4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 |
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/ | / | / | / | / | 11 | / | / | / | / |
Body band/ interspace ratio | 1.12 | 1.20 | 1.04 | 1.10 | 1.68 | 1.07 | 1.03 | 1.23 | 1.06 | 0.92 |
Precloacal groove | Deep | Deep | Deep | Deep | Deep | Absent | Absent | Absent | Absent | Absent |
Femoroprecloacal pores continuous | No | No | No | Yes | No | / | / | / | / | / |
Tuberculation | Weak | Weak | Weak | Weak | Weak | Weak | Weak | Weak | Weak | Weak |
Tubercles on ventral surface of forelimb | No | No | No | No | No | No | No | No | No | No |
Tubercles in gular region | No | No | No | No | No | No | No | No | No | No |
Ventrolateral fold tuberculate | No | No | No | No | No | No | No | No | No | No |
Dorsum bearing scattered pattern of white tubercles | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes |
Hatchlings/ juveniles with white tail tip | / | / | / | / | / | / | / | / | / | / |
Adult posterior caudal region white | / | No | / | / | / | No | / | No | / | / |
White caudal bands in adults immaculate | / | No | / | / | / | No | / | No | / | No |
Descriptive meristic (left/right) and color pattern characters of the referred specimens of
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Age |
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J |
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77.4 | 68.4 | 72.5 | 82.5 | 81.8 | 73.8 | 81.9 | 79.9 | 43.1 |
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12/12 | 13/13 | 12/12 | 12/13 | 14/14 | 14/14 | 13/13 | 13/14 | / |
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10/11 | 11/12 | 10/11 | 11/11 | 10/12 | 13/11 | 10/11 | 10/11 | / |
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38 | 38 | 32 | 41 | 41 | 41 | 42 | 40 | / |
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20 | 21 | 19 | 22 | 19 | 19 | 22 | 23 | / |
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34 | 40 | 34 | 37 | 32 | 37 | 38 | 37 | / |
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22/21 | 22/22 | 21/20 | 21/21 | 20/21 | 23/23 | 22/21 | 21/22 | / |
FFP in adult males | / | / | / | / | / | / | / | / | / |
/ | / | / | / | / | / | / | / | / | |
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4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 |
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11 | 12 | / | 11 | / | 10 | / | 11 | / |
Body band/ interspace ratio | 1.35 | 1.36 | 1.40 | 1.09 | 0.99 | 1.44 | 1.22 | 1.39 | / |
Precloacal groove | / | / | / | / | / | / | / | / | / |
Tuberculation | Weak | Weak | Weak | Weak | Weak | Weak | Weak | Weak | Weak |
Tubercles on ventral surface of forelimb | No | No | No | No | No | No | No | No | No |
Tubercles in gular region | No | No | No | No | No | No | No | No | No |
Ventrolateral fold tuberculate | No | No | No | No | No | No | No | No | No |
Dorsum bearing scattered pattern of white tubercles | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | No |
Hatchlings/juveniles with white tail tip | No | No | No | No | No | No | No | No | Yes |
Adult posterior caudal region white | / | / | / | / | / | / | / | / | / |
White caudal bands in adults immaculate | / | / | / | / | / | / | / | / | / |
In life, the juvenile (
Variation of
Habitat of
All specimens of
Two gravid females (
The specific epithet
Diagnostic characters of
Clade A | Clade B | ||||||||||||||||
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Sample size | 10 | 13 | 3 | 10 | 16 | 12 | 6 | 14 | 3 | 14 | 4 | 4 | 39 | 13 | 1 | 5 | 6 |
Maximum |
96.1 | 108.3 | 99.0 | 99.8 | 108.3 | 120.1 | 114.4 | 111.1 | 96.0 | 102.7 | 119.8 | 103.1 | 117.9 | 114.1 | 100.1 | 120.5 | 122.2 |
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12–15 | 10–12 | 12–14 | 9–12 | 10–12 | 9–12 | 8–12 | 9–13 | 9 or 10 | 9–11 | 10–12 | 10 or 11 | 9–12 | 11 | 10–12 | 9–12 | 9–13 |
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10–13 | 9–12 | 10–11 | 8–10 | 9–11 | 9–13 | 8 or 10 | 8–11 | 9 or 10 | 8–10 | 8–10 | 8–10 | 9–11 | 10 | 8–11 | 8–11 | 7–11 |
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32–47 | 40–57 | 35–36 | 34–44 | 30–50 | 37–45 | 36–44 | 31–42 | 31–34 | 33–43 | 38–43 | 36–41 | 36–42 | 31 | 34–38 | 39–48 | 34–49 |
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19–23 | 20–29 | 20–22 | 21–25 | 20–25 | 22–30 | 22–25 | 21–26 | 18–23 | 19–24 | 23–25 | 22–25 | 19–27 | 22–26 | 24 | 21–24 | 23–27 |
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32–40 | 31–46 | 36–39 | 38–43 | 30–43 | 32–40 | 31–39 | 36–40 | 29–33 | 26–33 | 31–36 | 32 or 33 | 17–28 | 29–34 | 41 | 31–40 | 33–36 |
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20–23 | 20–24 | 21–23 | 19–21 | 20–25 | 21–25 | 21–24 | 21–24 | 22–23 | 19–24 | 21–24 | 20–23 | 19–23 | 21–26 | 24 | 21–25 | 22–27 |
24–29 | 28–38 | 26–29 | 30 | 30–40 | 39–45 | 41–46 | 37–41 | 32–36 | 17–22 | 36 | 39–41 | 28–42 | 33–39 | 46 | 21 or 22 | 41–46 | |
11–15 | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | Absent | |
No. of body bands | 4 | 4 | 4 | 4 or 5 | 4 or 5 | 3(1) or 4 | 4 | 3(1) or 4 | 6 or 7 | 4 | 4 | 4 or 5 | 3–4 | 4 | 4 | 4 | 3 |
Body band/ interspace ratio | 0.92–1.68 | 0.98–2.07 | 0.75 | 0.75–1.00 | 0.86–2.00 | 1.00–2.00 | 1.00–1.25 | 1.00–1.25 | 0.82–1.10 | 1.00–1.25 | 1.00–1.50 | 0.50–1.25 | 0.95–1.74 | 0.75–1.25 | 1.75 | 1.00–1.25 | 2.00–2.75 |
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10–12 | 13 or 14 | >7 | 11–16 | 12–14 | 7–8 | 8 or 9 | 8–10 | 9–11 | 8–10 | 10 | 14 | 7–10 | 8–10 | 7 | 8–10 | 6–7 |
Precloacal groove in males | Deep | Deep | Deep | Deep | Deep | Deep | Deep | Deep | Shallow | Deep | Deep | Deep | Deep | Deep | Shallow | Deep | Deep |
Femoroprecloacal pores continuous | Both | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes | Yes |
Tuberculation | W | W | W | W | W | P | P | P | P | W | P | W | P | P | W | W | P |
Tubercles on ventral surface of forelimb | No | No | No | No | No | No | No | No | No | No | Yes | No | Yes | No | No | No | No |
Tubercles in gular region | No | No | No | No | No | No | No | No | No | No | No | No | Yes | No | No | No | No |
Ventrolateral fold tuberculate | No | No | No | No | No | No | Yes | No | No | No | No | No | Yes | No | No | No | No |
Dorsum bearing scattered pattern of white tubercles | Yes | Yes | Yes | No | No | No | No | No | No | No | No | No | No | No | No | No | No |
Hatchlings/juveniles with white tail tip | Yes | Yes | Yes | Yes | Yes | No | No | No | Yes | Yes | Yes | / | No | No | / | No | No |
Adult posterior caudal region white | No | No | No | No | No | No | No | No | Yes | Yes | No | No | No | No | No | No | No |
White caudal bands in adults immaculate | No | No | No | No | No | Yes | Yes | Yes | No | Yes | No | Yes | No | Yes | Yes | Yes | Yes |
Based on phylogenetic tree,
The discovery of
In addition, we found that the reported sampling localities of
The discovery of this new species suggests that undiscovered species of the
This research was supported by the Center of Excellence on Biodiversity (BDC), Office of Higher Education Commission (BDC-PG4-160022). Research protocol was approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project number ACKU61-SCI-006). KT was supported by a grant from the Faculty of Science, Kasetsart University (50th Anniversary of Faculty of Science). AR and AA were supported by Kasetsart University Research and Development Institute (KURDI), the Department of Zoology, and International SciKU Branding (ISB), Faculty of Science, Kasetsart University. NA was supported by Srinakharinwirot University Research Grant. The Department of National Parks, Wildlife and Plant Conservation, Thailand for providing permission to conduct the research in Tarutao National Park and Khao Banthat Wildlife Sanctuary. Kanchanapan Kamhang (Tarutao National Park) and Bamrungrat Ploydam (Khao Banthat Wildlife Sanctuary) for facilitating the fieldwork. L. Lee Grismer (La Sierra University), Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Siriporn Yodthong assisted with fieldwork. This paper is contribution number 950 of the Auburn University Museum of Natural History. We thank L. Lee Grismer and Evan S.H. Quah for providing their suggestions, which greatly improved the manuscript.