Corresponding author: Anchalee Aowphol (
Academic editor: T. Ziegler
The taxonomy and phylogeny of the
Termprayoon K, Rujirawan A, Grismer LL, Wood Jr PL, Aowphol A (2021) Taxonomic reassessment and phylogenetic placement of
The
During our field surveys, nine specimens of
During October 2017 and June 2019, field surveys were conducted at five localities in southern Thailand, including the type locality of
Specimens of
Species | Locality | Museum No. | GenBank Accession No. | Type of analysis | Reference |
---|---|---|---|---|---|
|
Unknow | NC 00155 |
|
A |
|
|
Pakistan, Baluchistan Province, Makran District, Gwadar | FMNH 247474 |
|
A |
|
|
Unknow | JB 28 |
|
A |
|
|
|
A |
|
||
|
|
A |
|
||
|
A |
|
|||
Laos, Khammouan Province, Nakai District | FMNH 255454 |
|
A |
|
|
FMNH 265806 |
|
A |
|
||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
BYU 53435 (holotype) |
|
A |
|
|
BYU 53436 (paratype) |
|
A |
|
||
BYU 53437 (paratype) |
|
A |
|
||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
BYU 51869 |
|
A |
|
|
BYU 51870 |
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A, B | |||
– | B | This study | |||
– | B | This study | |||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A, B | ||||
|
A |
|
|||
|
A |
|
|||
– | B | This study | |||
|
A |
|
|||
|
A |
|
|||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
|
A |
|
|||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
|
A, B | This study | |||
– | B | This study | |||
– | B | This study | |||
– | B | This study | |||
|
A, B | This study | |||
– | B | This study | |||
– | B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A, B | This study | |||
|
A | This study | |||
PSUZC-RT 2010.58 | – | B | This study | ||
– | B | This study | |||
|
|
A |
|
||
|
A |
|
|||
|
A, B | ||||
|
A, B | ||||
|
A, B | ||||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
A, B | ||||
– | B | This study | |||
– | B | This study | |||
|
|
A |
|
||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
|||
|
A |
|
|||
|
|
A |
|
||
|
A |
|
|||
|
A |
|
Map illustrating the known geographic distribution of
Total genomic DNA was extracted from 95% ethanol-preserved muscle or liver tissue using a NucleoSpin Tissue Kit (Macherey-Nagel GmbH & Co. KG, Germany). Mitochondrial NADH dehydrogenase subunit 2 (
Phylogenetic relationships were inferred using two model based approaches, Bayesian Inference (
Morphological and meristic characters were modified from the previous studies of
Additional scale counts and non-meristic characters evaluated were the number of supralabial and infralabial scales counted from the largest scale immediately posterior to the dorsal inflection of the posterior portion of the upper jaw to the rostral and mental scales, respectively; the number of paravertebral tubercles between limb insertions counted in a straight line immediately left of the vertebral column; the number of longitudinal rows of body tubercles counted transversely across the center of the dorsum from one ventrolateral fold to the other; the number of longitudinal rows of ventral scales counted transversely across the center of the abdomen from one ventrolateral fold to the other; the presence or absence of tubercles on the ventral surface of the forearm; the presence or absence of tubercles in the gular region, throat, and lateral margins of the abdomen; the number of subdigital lamellae beneath the fourth toe counted from the base of the first phalanx to the claw; the total number of precloacal and femoral pores (i.e., the contiguous rows of femoral and precloacal scales bearing pores combined as a single meristic referred to as the femoroprecloacal pores); the presence or absence of a precloacal depression or groove; the degree of body tuberculation, weak tuberculation referring to dorsal body tubercles that are low and rounded whereas prominent tuberculation refers to tubercles that are raised and keeled; the width of the dark body bands relative to the width of the interspace between the bands; number of dark caudal bands on the original tail; the white caudal bands of adults immaculate or infused with dark pigment; and whether or not the posterior portion of the original tail in hatchlings and juveniles less than 50 mm
All statistical analyses were performed using the base statistical software in RStudio v. 1.2.1335 (
Principal component analysis (
For univariate analyses, all transformed mensural characters were tested for normality using the Shapiro-Wilk Test. Equality of variances was tested using F-tests. Morphological differences of both males and females between
The aligned matrix contained 1,453 bp of
Percentage uncorrected pairwise sequence divergence (
Species |
|
1 | 2 | 3 | 4 | |
---|---|---|---|---|---|---|
1 |
|
9 |
|
|||
2 |
|
3 | 7.42 (6.64–8.38) | |||
3 |
|
27 | 8.93 (7.47–10.48) | 8.08 (6.64–8.38) | ||
4 |
|
9 | 9.41 (8.71–10.29) | 8.79 (8.45–8.95) | 2.63 (1.45–4.20) |
Reconstructed phylogenetic relationships of the
A total of 45 preserved specimens from three species groups (
Summary statistics and factor loadings of the principal component analysis from morphological characters for males and females
Characters | Males | Females | ||||
---|---|---|---|---|---|---|
|
|
|
|
|
|
|
|
0.067 | -0.193 | -0.291 | 0.080 | -0.043 | -0.047 |
|
0.558 | 0.589 | -0.029 | -0.823 | 0.036 | 0.055 |
|
0.586 | -0.425 | 0.023 | -0.040 | 0.507 | -0.474 |
|
0.467 | -0.690 | 0.027 | -0.055 | 0.452 | -0.752 |
|
-0.185 | -0.442 | 0.415 | 0.685 | -0.284 | -0.283 |
|
0.784 | 0.141 | -0.033 | 0.384 | 0.730 | 0.436 |
|
0.439 | 0.569 | -0.144 | 0.232 | 0.405 | -0.163 |
|
0.577 | 0.539 | -0.221 | 0.516 | -0.070 | 0.040 |
|
0.585 | -0.173 | 0.374 | -0.364 | 0.777 | 0.337 |
|
0.484 | 0.314 | -0.580 | 0.581 | 0.338 | -0.056 |
|
0.858 | -0.053 | -0.017 | 0.669 | 0.260 | 0.455 |
|
0.774 | -0.289 | 0.004 | 0.726 | -0.087 | 0.133 |
|
0.147 | 0.662 | 0.003 | 0.602 | 0.117 | 0.174 |
|
0.351 | -0.097 | -0.360 | 0.141 | 0.190 | 0.714 |
INadj | 0.527 | 0.224 | -0.184 | -0.216 | 0.150 | -0.473 |
Supralabials | -0.217 | 0.622 | 0.270 | -0.070 | -0.745 | 0.307 |
Infralabials | 0.218 | 0.627 | 0.402 | -0.258 | -0.690 | 0.340 |
Paravertebral tubercles | -0.157 | 0.472 | 0.409 | -0.287 | -0.196 | -0.168 |
Longitudinal tubercles | 0.363 | 0.215 | 0.642 | -0.133 | 0.215 | 0.464 |
Ventral scales | 0.761 | -0.297 | 0.338 | -0.665 | 0.497 | 0.230 |
4th toe lamellae | 0.439 | -0.324 | -0.217 | -0.864 | 0.095 | 0.283 |
Femoroprecloacal pores | 0.512 | -0.196 | 0.468 | / | / | / |
Eigenvalue | 5.671 | 3.864 | 2.164 | 4.859 | 3.484 | 2.784 |
Percentage of variance | 25.776 | 17.562 | 9.834 | 23.136 | 16.592 | 13.258 |
Cumulative proportion | 25.776 | 43.338 | 53.172 | 23.136 | 39.728 | 52.986 |
Plots for the first two principal components of morphological characters from
Along the first two PC plots, the
Summary univariate statistics of morphological characters of adult males and females are shown in Table
Comparisons of fifteen morphological characters between
Characters | Males | Females | ||||||
---|---|---|---|---|---|---|---|---|
|
|
|
|
|
|
|||
|
|
|||||||
|
105.8 ± 8.9 | 105.3 ± 10.0 | 0.112 | 0.9122 | 103.7 ± 10.1 | 104.4 ± 16.0 | -0.098 | 0.9237 |
(88.9–117.9) | (93.2–115.3) | (84.1–115.7) | (84.8–117.6) | |||||
|
9.5 ± 0.9 | 9.6 ± 1.2 | -0.061 | 0.9523 | 8.1 ± 1.4 | 8.2 ± 2.1 | -0.352 | 0.7307 |
(8.0–11.6) | (7.9–10.9) | (5.9–10.3) | (6.2–10.5) | |||||
|
17.2 ± 1.5 | 17.2 ± 2.0 | -0.098 | 0.9229 | 16.8 ± 1.6 | 16.8 ± 2.7 | 0.048 | 0.9626 |
(14.2–18.9) | (14.4–19.0) | (13.6–18.3) | (13.3–19.3) | |||||
|
20.3 ± 1.6 | 20.6 ± 2.6 | -0.631 | 0.5348 | 19.9 ± 1.8 | 20.4 ± 3.2 | -2.166 | 0.0495* |
(17.3–22.8) | (17.4–19.0) | (16.3–21.9) | (16.5–23.2) | |||||
|
50.4 ± 4.6 | 51.5 ± 5.9 | -1.034 | 0.3125 | 50.7 ± 4.5 | 53.4 ± 7.9 | 4b | 0.0176* |
(41.3–58.6) | (44.7–57.4) | (42.0–56.0) | (43.4–60.2) | |||||
|
29.4 ± 2.2 | 29.2 ± 2.7 | 0.590 | 0.5613 | 28.5 ± 3.1 | 28.4 ± 4.4 | 0.575a | 0.5758 |
(24.6–33.3) | (25.6–31.7) | (22.8–32.3) | (23.1–32.1) | |||||
|
20.0 ± 1.8 | 19.8 ± 2.7 | 0.416 | 0.6816 | 19.0 ± 1.9 | 18.7 ± 2.6 | 0.815 | 0.4297 |
(16.7–22.9) | (16.3–22.5) | (15.6–21.0) | (15.8–21.4) | |||||
|
12.0 ± 1.3 | 12.2 ± 1.9 | -0.840 | 0.4100 | 11.3 ± 1.3 | 11.5 ± 2.0 | -1.054 | 0.3109 |
(9.7–14.1) | (9.9–14.2) | (9.0–13.4) | (9.1–13.5) | |||||
|
6.9 ± 0.6 | 6.7 ± 0.5 | 1.397 | 0.1765 | 6.7 ± 0.7 | 6.5 ± 1.1 | 1.253 | 0.2325 |
(5.8–7.9) | (5.6–7.0) | (5.6–7.6) | (7.4–9.6) | |||||
|
8.6 ± 1.0 | 8.6 ± 1.0 | 62b | 0.6261 | 8.5 ± 0.8 | 8.6 ± 1.1 | -0.106 | 0.9171 |
(6.5–9.8) | (7.1–9.4) | (7.0–9.4) | (7.4–9.6) | |||||
|
11.7 ± 1.0 | 11.7 ± 1.1 | 56b | 0.9225 | 11.5 ± 1.2 | 11.4 ± 1.8 | 0.271a | 0.7912 |
(10.0–13.6) | (10.3–12.9) | (9.3–13.1) | (9.2–12.9) | |||||
|
8.7 ± 0.7 | 8.7 ± 0.7 | 0.090 | 0.9288 | 8.6 ± 0.9 | 8.5 ± 1.3 | 0.610 | 0.5521 |
(7.3–9.8) | (7.6–9.5) | (6.8–9.8) | (6.8–9.6) | |||||
|
5.1 ± 0.6 | 4.8 ± 0.6 | 2.394 | 0.0256* | 4.8 ± 0.8 | 4.6 ± 0.9 | 0.835 | 0.4186 |
(4.0–6.3) | (4.1–5.5) | (3.4–5.7) | (3.6–5.5) | |||||
|
2.3 ± 0.3 | 2.2 ± 0.4 | 0.504 | 0.6192 | 2.4 ± 0.4 | 2.2 ± 0.5 | 1.125 | 0.281 |
(1.7–2.8) | (1.4–2.6) | (1.6–3.0) | (1.5–2.5) | |||||
IN | 2.2 ± 0.3 | 2.3 ± 0.3 | -0.352 | 0.7282 | 2.0 ± 0.5 | 2.3 ± 0.3 | -1.785a | 0.0129* |
(1.7–2.7) | (1.7–2.5) | (1.3–2.9) | (1.9–2.6) |
Summarized diagnostic characters of
|
|
|
|
|
---|---|---|---|---|
(Grismer and Ahmad, 2008) | ( |
This study | This study | |
Supralabials | 10–12 | 12–13 | 9–12 | 9–13 |
Infralabials | 8–11 | 9–10 | 7–11 | 7–10 |
Tuberculation | Prominent | Prominent | Prominent | Prominent |
Tubercles on ventral surface of forelimbs | Yes | Yes | Yes | Yes |
Tubercles in gular region | Yes | Yes | Yes | Yes |
Ventrolateral fold tuberculate | Yes | Yes | Yes | Yes |
Paravertebral tubercles | 40–47 | 40–43 | 34–49 | 39–45 |
Longitudinal rows of tubercles | 22–26 | 23–24 | 19–27 | 20–24 |
Ventral scales | 19–22 | 22–24 | 17–28 | 20–24 |
4th toe lamellae | 21–23 | 19 | 19–23 | 19–21 |
Femoroprecloacal pores | 35–37 | 33–36 | 28–42 | 31–33 |
Precloacal groove present in females | No | Yes | No | No |
Precloacal depression in males | No | No | Deep | Deep |
No. of body bands | 4 | 3 (3.5 one individual) | 3–4 | 3–4 (3+1 incomplete band) |
Body band/interspace ratio | / | / | 0.95–1.74 | 1.02–1.50 |
Dorsum bearing scattered pattern of white tubercles | No | No | No | No |
Hatchlings/juveniles with white tail tip | No | No | No | No |
Dark caudal bands on original tail | / | 8 | 7–10 | 7–8 |
White caudal bands in adults immaculate | / | / | No | No |
Maximum |
120.0 | 114.7 | 117.87 | 117.61 |
Sample size | 5 | 3 | 29 | 10 |
The phylogenetic analyses recovered
Morphological measurement (mm), meristic and non-meristic data from males and females of
Characters | Adult males ( |
Adult females ( |
All ( |
|||
---|---|---|---|---|---|---|
Mean ± SD | (Min–Max) | Mean ± SD | (Min–Max) | Mean ± SD | (Min–Max) | |
|
105.7 ± 9.0 | (88.9–117.9) | 103.9 ± 11.3 | (84.1–117.6) | 105.0 ± 9.8 | (84.1–117.9) |
|
9.5 ± 1.0 | (7.9–11.6) | 8.2 ± 1.5 | (5.9–10.5) | 9.0 ± 1.4 | (5.9–11.6) |
|
17.2 ± 1.6 | (14.2–19.0) | 16.8 ± 1.8 | (13.3–19.3) | 17.1 ± 1.7 | (13.3–19.3) |
|
20.4 ± 1.9 | (17.3–23.5) | 20.0 ± 2.1 | (16.3–19.3) | 20.2 ± 2.0 | (16.3–23.5) |
|
50.7 ± 4.9 | (41.3–58.6) | 51.4 ± 5.4 | (42.0–60.2) | 51.0 ± 5.0 | (41.3–60.2) |
|
29.4 ± 2.3 | (24.6–33.3) | 28.5 ± 3.3 | (22.8–32.3) | 29.0 ± 2.7 | (22.8–33.3) |
|
20.0 ± 2.0 | (16.3–22.9) | 18.9 ± 2.0 | (15.6–21.4) | 19.6 ± 2.0 | (15.6–22.9) |
|
12.1 ± 1.4 | (9.7–14.2) | 11.3 ± 1.4 | (9.0–13.5) | 11.8 ± 1.4 | (9.0–14.2) |
|
6.8 ± 0.6 | (5.6–7.9) | 6.7 ± 0.8 | (5.2–7.5) | 6.8 ± 0.7 | (5.2–7.9) |
|
8.6 ± 1.0 | (6.5–9.8) | 8.5 ± 0.9 | (7.0–9.6) | 8.6 ± 0.7 | (5.2–7.9) |
|
11.7 ± 1.0 | (10.0–13.6) | 11.5 ± 1.3 | (9.2–13.1) | 11.6 ± 1.1 | (9.2–13.6) |
|
8.7 ± 0.7 | (7.3–9.8) | 8.6 ± 1.0 | (6.8–9.8) | 8.6 ± 0.8 | (6.8–9.8) |
|
5.0 ± 0.6 | (4.0–6.3) | 4.8 ± 0.8 | (3.4–5.7) | 4.8 ± 0.7 | (3.4–6.3) |
|
2.3 ± 0.3 | (1.4–2.8) | 2.3 ± 0.4 | (1.5–3.0) | 2.3 ± 0.4 | (1.4–3.0) |
IN | 2.2 ± 0.5 | (1.7–2.7) | 2.1 ± 0.5 | (1.3–2.9) | 2.2 ± 0.4 | (1.3–2.9) |
0.28 ± 0.01 | (0.27–0.30) | 0.27 ± 0.01 | (0.26–0.29) | 0.28 ± 0.01 | (0.26–0.30) | |
0.68 ± 0.03 | (0.62–0.74) | 0.67 ± 0.02 | (0.62–0.70) | 0.67 ± 0.03 | (0.62–0.74) | |
0.41 ± 0.02 | (0.37–0.45) | 0.40 ± 0.01 | (0.38–0.42) | 0.41 ± 0.02 | (0.37–0.45) | |
0.40 ± 0.01 | (0.37–0.41) | 0.40 ± 0.00 | (0.39–0.41) | 0.40 ± 0.01 | (0.37–0.41) | |
0.23 ± 0.01 | (0.21–0.27) | 0.23 ± 0.01 | (0.22–0.25) | 0.23 ± 0.01 | (0.21–0.27) | |
0.08 ± 0.01 | (0.05–0.10) | 0.08 ± 0.01 | (0.05–0.10) | 0.08 ± 0.01 | (0.05–0.10) | |
0.48 ± 0.02 | (0.43–0.51) | 0.50 ± 0.01 | (0.47–0.52) | 0.49 ± 0.02 | (0.43–0.52) | |
0.16 ± 0.00 | (0.15–0.17) | 0.16 ± 0.00 | (0.16–0.17) | 0.16 ± 0.00 | (0.15–0.17) | |
0.19 ± 0.01 | (0.18–0.21) | 0.19 ± 0.00 | (0.18–0.20) | 0.19 ± 0.01 | (0.18–0.21) | |
1.29 ± 0.04 | (1.23–1.35) | 1.27 ± 0.04 | (1.24–1.34) | 1.28 ± 0.04 | (1.23–1.35) | |
Supralabials | 9–13 | 9–12 | 9–13 | |||
Infralabials | 7–11 | 7–11 | 7–11 | |||
Tuberculation | Prominent | Prominent | Prominent | |||
Tubercles on ventral surface of forelimbs | Yes | Yes | Yes | |||
Tubercles in gular region | Yes | Yes | Yes | |||
Ventrolateral fold tuberculate | Yes | Yes | Yes | |||
Paravertebral tubercles | 37–49 | 34–47 | 34–49 | |||
Longitudinal rows of tubercles | 19–27 | 19–26 | 19–27 | |||
Ventral scales | 17–28 | 19–26 | 17–28 | |||
4th toe lamellae | 19–23 | 19–23 | 19–23 | |||
Femoroprecloacal pores | 28–42 | No | 28–42 | |||
Precloacal depression | Yes | No | Only in males | |||
No. of body bands | 3 or 4 | 3 or 4 | 3 or 4 | |||
Body band/interspace ratio | 0.95–1.75 | 1.03–1.62 | 0.95–1.74 | |||
Dark caudal bands on original tail | 7–9 | 7–10 | 7–10 |
Body relatively short (
Forelimbs moderate in stature, relatively short (
Original tail (
Male holotype of
Adult male
Adult male
Variation in dorsal body band pattern of
Color in life of
Precloacal region in female specimens of
Based on specimens in Thailand, all individuals were found in similar habitat type, lowland forest habitat along granitic rock streams and surrounding areas (elevation 7–186 m asl) during a night survey (1900–2200; Fig.
Habitats of
In Thailand,
Diagnostic characters of
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Supralabials | 9–12 | 9–13 | 8–12 | 9 or 10 | 10 or 11 | 9–11 | 11 | 10–12 | 9–13 | 10–12 | 12–14 | 9–12 | 9–12 | 10–12 | 9–12 | 10–12 |
Infralabials | 9–13 | 8–11 | 8 or 10 | 9 or 10 | 8–10 | 8–10 | 10 | 8–11 | 7–11 | 9–12 | 10–11 | 8–11 | 8–10 | 9–11 | 9–11 | 8–10 |
Tuberculation | P | P | P | P | W | P | W | P | P | W | W | W | W | W | P | W |
Tubercles on ventral surface of forelimbs | No | No | No | No | No | No | No | No | Yes | No | No | No | No | No | Yes | No |
Tubercles in gular region | No | No | No | No | No | No | No | No | Yes | No | No | No | No | No | No | No |
Ventrolateral fold tuberculate | No | No | Yes | No | No | No | No | No | Yes | No | No | No | No | No | No | No |
Paravertebral tubercles | 37–45 | 31–42 | 36–44 | 31–34 | 36–41 | 33–43 | 31 | 34–38 | 34–49 | 40–57 | 35–36 | 39–48 | 34–44 | 30–50 | 36–42 | 38–43 |
Longitudinal rows of tubercles | 22–30 | 21–26 | 22–25 | 18–23 | 22–25 | 22–26 | 24 | 23–27 | 19–27 | 20–29 | 20–22 | 19–24 | 21–25 | 20–24 | 23–25 | 21–24 |
Ventral scales | 32–40 | 36–40 | 31–39 | 29–33 | 32 or 33 | 29–34 | 41 | 33–36 | 17–28 | 31–46 | 36–39 | 26–33 | 38–43 | 31–43 | 31–36 | 31–40 |
4th toe lamellae | 21–25 | 21–24 | 21–24 | 22–23 | 20–23 | 21–26 | 24 | 22–27 | 19–23 | 20–24 | 21–23 | 19–24 | 19–21 | 20–25 | 21–24 | 21–25 |
Femoroprecloacal pores | 39–45 | 37–41 | 41–46 | 32–36 | 39–41 | 33–39 | 46 | 41–46 | 28–42 | 31–38 | 26–29 | 17–22 | 30 | 30–36 | 36 | 21 or 22 |
Precloacal groove in males | Deep | Deep | Deep | Shallow | Deep | Deep | Shallow | Deep | Deep | Deep | Deep | Deep | Deep | Deep | Deep | Deep |
No. of body bands | 3(1) or 4 | 3(1) or 4 | 4 | 6 or 7 | 4 or 5 | 4 | 4 | 3 | 3–4 | 4 | 4 | 4 | 4 or 5 | 4 or 5 | 4 | 4 |
Body band/interspace ratio | 1.00–2.00 | 1.00–1.25 | 1.00–1.25 | 0.82–1.10 | 0.50–1.25 | 0.75–1.25 | 1.75 | 2.00–2.75 | 0.95–1.74 | 1.00–2.00 | 0.75 | 1.00–1.25 | 0.75–1.00 | 1.00–2.00 | 1.00–1.50 | 1.00–1.25 |
Dorsum bearing scattered pattern of white tubercles | No | No | No | No | No | No | No | No | No | Yes | Yes | No | No | No | No | No |
Hatchlings/juveniles with white tail tip | No | No | No | Yes | / | No | / | No | No | Yes | Yes | Yes | Yes | Yes | Yes | No |
Adult posterior caudal region white | No | No | No | Yes | No | No | No | No | No | No | No | Yes | No | No | No | No |
Dark caudal bands on original tail | 7–8 | 8–10 | 8 or 9 | 9–11 | 14 | 8–10 | 7 | 6–7 | 7–10 | 13 or 14 | >7 | 8–10 | 11–16 | 12–14 | 10 | 8–10 |
White caudal bands in adults immaculate | Yes | Yes | Yes | No | Yes | Yes | Yes | Yes | No | No | No | Yes | No | No | No | Yes |
Maximum |
120.10 | 111.1 | 114.40 | 96.00 | 103.1 | 114.1 | 100.1 | 122.2 | 117.9 | 108.3 | 99.00 | 102.7 | 99.8 | 103.5 | 119.8 | 120.5 |
Sample size | 12 | 14 | 6 | 3 | 4 | 13 | 1 | 6 | 39 | 11 | 3 | 14 | 10 | 13 | 4 | 5 |
Based on molecular data,
Within the
Evidence from both overlapping ranges of morphology and relatively low sequence divergence indicate that
This work was supported by grants from the Thailand Research Fund (DBG6080010) and the Center of Excellence on Biodiversity (BDC), Office of Higher Education Commission (BDC-PG4-160022). AA was supported by the Department of Zoology and International SciKU Branding (ISB), Faculty of Science, Kasetsart University and KT was supported by a grant from the Faculty of Science, Kasetsart University (50th Anniversary of Faculty of Science). This research was approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project number ACKU61-SCI-006). The Department of National Parks, Wildlife and Plant Conservation, Thailand, granted permission to conduct the research. We thank Piyawat Sukon (superintendent of Khao Phra Thaeo Non-hunting Area), Kanchanapan Kamhang (superintendent of Tarutao National Park), and Chai Suvannachat (superintendent of Ton Nga Chang Wildlife Sanctuary) for facilitating the fieldwork. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Siriporn Yodthong, Natee Ampai, Akrachai Aksornneam and Piyawan Puanprapai assisted with fieldwork. This paper is contribution number 919 of the Auburn University Museum of Natural History.