Corresponding author: Anchalee Aowphol (
Academic editor: A. Bauer
The rock-dwelling gecko genus
Ampai N, Wood Jr PL, Stuart BL, Aowphol A (2020) Integrative taxonomy of the rock-dwelling gecko
The rock-dwelling gecko genus
Southeast Asian
The
More than half of Southeast Asian
During fieldwork in October 2016 and January 2019, we collected specimens of the
Specimens of
The following morphometric measurements were taken by the first author on the left side of preserved specimens to the nearest 0.1 mm using digital calipers under a Nikon SMZ 745 dissecting microscope. Morphological measurements were taken only from adult individuals as determined by the presence of secondary sexual characteristics including the presence of hemipenes or pore-bearing precloacal scales in males, and the presence of calcium glands or eggs in females. Sixteen morphological measurements were taken following
Meristic characters of scales and qualitative observations of other structures were made through a Nikon SMZ 745 dissecting microscope. The external observations of meristic characters were taken following
Statistical analyses were used to compare differences in size and shape in the
Multivariate analyses were performed using the base statistical software in RStudio v. 1.2.1335 (
Genomic DNA was extracted from liver tissue of five individuals of
Specimens used in this study, including locality, collection numbers and Genbank accession numbers. Voucher abbreviations are as follows:
Species | Locality | Collection number | GenBank accession number | Reference |
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Cambodia, Kampot |
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Myanmar, Mon State, |
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Kyait Hti Yo Wildlife Sanctuary | ||||
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Thailand, Satun Province, |
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Mueang Satun District, |
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Adang Island |
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Thailand, Satun Province, |
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Mueang Satun District, Rawi Island |
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Malaysia, Penang, Pulau Pinang |
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Malaysia, Terengganu, |
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Gunung Lawit |
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Vietnam, Kien Giang Province, Hon Dat Hill |
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Malaysia, Perlis, Kuala Perlis |
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Malaysia, Perlis, Gua Kelam |
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Vietnam, Ca Mau Province, |
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Con Dao Archipelago |
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Vietnam, Kien Giang Province, Hon Tre Island |
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Thailand, Nakhon Si Thammarat Province, Thum Thong Panra |
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Cambodia, Pursat Province, Phnom Dalai |
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Malaysia, Perak, Lenggong |
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Malaysia, Pahang, Bukit Hangus |
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Malaysia, Kedah, Gunung Jeri |
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Thailand, Kanchanaburi Province, Sai Yok National Park |
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Malaysia, Kelantan, Gunung Reng |
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Malaysia, Perlis, Perlis State Park |
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Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai Pak Waterfall |
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This study | ||
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This study | |||
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This study | |||
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This study | |||
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This study | |||
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Thailand, Prachuap Khiri Khan Province, |
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Kui Buri District, Wat Khao Daeng |
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Malaysia, Kedah, Pulau Langkawi, Gunung Raya |
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Malaysia, Perak, Bukit Larut |
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Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi |
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Malaysia, Perak, Belum-Temengor, Sungai Enam |
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Cambodia, Pursat Province, O’Lakmeas |
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Thailand, Trang Province, |
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Thum Khao Ting |
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Vietnam, An Giang Province, Nui Cam Hill |
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Thailand, Satun Province, Phuphaphet Cave |
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Malaysia, Perlis, Perlis State Park |
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Malaysia, Terengganu, Pulau Perhentian Besar |
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Thailand, Phangnga Province, |
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Mueang Phangnga District, Khao Chang, Phung Chang Cave |
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Thailand, Prachaup Khiri Khan Province, Thap Sakae |
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Malaysia, Kedah, Pulau Langkawi, Gunung Raya |
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Thailand, Chumpon Province, Pathio District |
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Thailand, Satun Province, |
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Mueang Satun District, | ZMKUR 00763 |
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Tarutao Island |
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Thailand, Surat Thani Province, |
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Tha Chana District, |
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Tham Khao Sonk Hill |
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Vietnam, An Giang Province, |
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Tuc Dup Hill |
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Thailand, Ranong Province, Suk Saran District, Naka |
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Homologous sequences of 69
Phylogenies were reconstructed with the maximum likelihood (
Phylogenies were also reconstructed with Bayesian Inference (
The most likely tree in the
The
The
Results of principal component analysis (
Pairwise matrix of significant differences (Tukey’s pairwise;
Species |
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Summary of eigenvalues, percentage of variance, standard deviation, cumulative proportion, and factor loadings from the principal components (
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5.083 | 3.855 | 2.564 | 1.308 | 0.520 | 0.385 | 0.347 | 0.237 | 0.204 | 0.173 | 0.131 | 0.084 | 0.050 | 0.037 | 0.021 |
% |
33.884 | 25.697 | 17.097 | 8.721 | 3.466 | 2.565 | 2.316 | 1.578 | 1.362 | 1.156 | 0.875 | 0.558 | 0.336 | 0.247 | 0.142 |
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2.254 | 1.963 | 1.601 | 1.144 | 0.721 | 0.620 | 0.589 | 0.486 | 0.452 | 0.417 | 0.362 | 0.289 | 0.224 | 0.193 | 0.146 |
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0.339 | 0.596 | 0.767 | 0.854 | 0.889 | 0.914 | 0.937 | 0.953 | 0.967 | 0.978 | 0.987 | 0.993 | 0.996 | 0.999 | 1.000 |
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0.254 | 0.298 | 0.093 | -0.016 | 0.156 | 0.841 | 0.112 | -0.264 | 0.077 | -0.008 | 0.126 | -0.034 | 0.002 | 0.039 | -0.003 |
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0.126 | -0.028 | -0.226 | 0.142 | -0.163 | 0.093 | 0.487 | 0.221 | -0.048 | 0.583 | -0.289 | -0.078 | -0.050 | 0.119 |
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0.085 | 0.117 |
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0.292 | -0.046 | -0.103 | 0.343 | 0.186 | -0.172 | -0.080 | 0.046 | -0.006 | 0.176 | 0.192 | -0.583 |
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0.197 | 0.277 | 0.238 |
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0.040 | -0.064 | 0.011 | 0.160 | -0.020 | -0.103 | -0.325 | -0.086 | -0.029 | -0.473 | 0.448 |
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-0.158 | - |
-0.172 |
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-0.416 | 0.387 | -0.024 | 0.461 | 0.140 | -0.163 | 0.039 | -0.013 | -0.322 | 0.110 | -0.037 |
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0.086 |
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-0.181 |
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0.068 | -0.232 | -0.321 | -0.231 | 0.176 | -0.088 | 0.012 | -0.203 | -0.234 | 0.555 | -0.033 |
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-0.179 | 0.206 | - |
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0.115 | -0.051 | 0.113 | -0.072 | 0.133 | 0.353 | 0.289 | 0.175 | -0.008 | -0.452 | -0.363 |
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0.168 | -0.007 | - |
0.026 | -0.185 | -0.063 | 0.502 | -0.196 | -0.278 | -0.409 | -0.078 | -0.317 | 0.155 | -0.029 | -0.005 |
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0.296 | -0.294 | 0.176 | 0.082 | -0.091 | -0.161 | 0.398 | -0.359 | 0.022 | 0.254 | 0.112 | 0.158 | -0.586 | 0.036 | 0.142 |
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0.194 |
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-0.205 | -0.088 | -0.199 | 0.009 | -0.003 | 0.278 | -0.505 | 0.148 | 0.022 | 0.551 | -0.067 | 0.202 | 0.145 |
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-0.042 | -0.010 | -0.143 | -0.082 | -0.050 | -0.395 | -0.110 | 0.026 | -0.471 | -0.055 | 0.243 | -0.239 | -0.342 | -0.417 |
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-0.058 | 0.030 | 0.059 | -0.639 | -0.009 | -0.244 | -0.144 | 0.093 | 0.402 | 0.064 | -0.187 | 0.394 | -0.045 | -0.009 |
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-0.090 | -0.240 | -0.057 | 0.216 | -0.003 | 0.211 | 0.244 | 0.459 | 0.191 | -0.553 | 0.174 | 0.126 | 0.170 | -0.141 |
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0.178 | - |
-0.040 |
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0.248 | -0.020 | -0.072 | -0.104 | 0.028 | -0.239 | 0.335 | 0.407 | 0.441 | 0.143 | 0.232 |
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0.244 | - |
-0.119 | 0.149 | 0.395 | 0.111 | -0.263 | 0.121 | -0.542 | 0.302 | -0.081 | -0.341 | -0.102 | -0.005 | -0.150 |
The aligned dataset contained 1,251 characters of 69 individuals of
The Lan Saka samples represented a well-supported clade (100
Mean (min-max) uncorrected
No. | Species | N | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 |
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1 | 5 |
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2 |
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5 | 17.78 |
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(17.02–18.72) |
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3 |
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1 | 16.98 | 11.40 |
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(16.81–17.23) | (10.85–11.91) | ||||||||||||
4 |
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3 | 27.45 | 9.36 | 11.81 |
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(27.23–27.87) | (8.30–10.21) | (11.49–12.13) |
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5 |
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2 | 18.79 | 10.19 | 11.38 | 11.17 |
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(18.51–19.15) | (9.57–10.85) | (11.27–11.49) | (10.85–11.49) |
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6 |
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2 | 17.64 | 25.83 | 24.40 | 27.77 | 25.00 |
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(17.45–17.87) | (25.74–25.96) | (24.26–24.68) | (27.66–27.87) | (24.89–25.11) | |||||||||
7 |
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2 | 25.91 | 8.92 | 11.77 | 9.01 | 8.90 | 28.16 |
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(25.74–26.17) | (8.51–9.57) | (11.70–11.91) | (8.72–9.36) | (8.72–9.15) | (28.09–28.30) |
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8 |
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3 | 15.77 | 24.26 | 24.04 | 26.60 | 25.21 | 12.34 | 26.88 |
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(15.53–16.17) | (24.04–24.47) | (24.04–24.04) | (25.96–27.23) | (25.11–25.32) | (12.34–12.34) | (26.81–27.02) | |||||||
9 |
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2 | 27.74 | 25.50 | 24.40 | 28.30 | 26.13 | 13.35 | 27.66 | 14.47 |
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(27.66–27.87) | (25.10–25.96) | (24.26–24.68) | (27.23–28.94) | (25.74–26.81) | (13.19–14.26) | (27.45–28.09) | (14.26–14.89) |
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10 |
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3 | 27.53 | 25.23 | 25.53 | 26.38 | 25.00 | 19.36 | 25.60 | 21.06 | 21.13 |
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(27.23–28.09) | (24.04–26.17) | (25.53–25.53) | (26.38–26.38) | (25.00–25.00) | (19.36–19.36) | (25.53–25.74) | (21.06–21.06) | (21.06–21.28) | |||||
11 |
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1 | 25.62 | 26.00 | 26.17 | 28.19 | 23.72 | 19.36 | 27.52 | 20.64 | 20.99 | 16.95 |
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(25.53–25.74) | (25.74–26.38) | (26.17–26.17) | (27.87–28.51) | (23.62–23.83) | (19.36–19.36) | (27.45–27.66) | (20.64–20.64) | (20.64–21.70) | (16.81–17.02) |
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Male holotype (
Male holotype (
Male holotype (
Meristic character state and color pattern of species in the
Characters/Species |
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Sample size | 19 | 15 | 25 | 5 | 3 | 8 | 2 | 5 | 8 | 12 | 6 | 3 |
Maximum |
41.8 | 44.9 | 40.9 | 43.5 | 37.8 | 41.3 | 42.0 | 49.6 | 47.0 | 39.7 | 39.0 | 44.7 |
Supralabial scales | 8–9 | 10 | 8–10 | 7–10 | 8–9 | 8–9 | 10 | 8 | 8–9 | 8–9 | 10–11 | 8–9 |
Infralabial scales | 8–9 | 9 | 8 | 6–9 | 7–8 | 7–8 | 10 | 7–8 | 7–8 | 6–8 | 9–11 | 7–9 |
Ventral scales keeled (1) or smooth (0) | 1 | 1 | 1 | 0 | w,0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 |
No. of pore-bearing precloacal scales | 4–7 | 6–8 | 6–8 | 5–8 | 6–7 | 3–6 | 4 | 0 | 3–6 | 0 | 0 | 4 |
Precloacal scales pore-bearing continuous (1) or separated (0) | 0 | 0 | 0 | 1 | 1 | 0 | 1 | – | 0 | – | – | 0 |
Precloacal pores elongate (1) or round (0) | 0 | 0 | 0 | 0 | 1 | 0 | 0 | – | 0 | – | – | 0 |
No. of paravertebral tubercles | 19–21 | 23–25 | 22–25 | 18–24 | 19–24 | 22–29 | 22 | 24–27 | 25–27 | 19–25 | 15–19 | 25–29 |
Paravertebral tubercles linearly arranged (1) or more random (0) | 1 | 0 | 0 | w,0 | w | w,0 | 1 | w | 0 | 0 | 1 | 0 |
Tubercles present (1) or absent (0) on lower flanks | 1 | 0 | 1 | 1 | 1 | w,1 | 0 | 1 | 1 | 1 | 1 | 1 |
No. of 4th toe lamellae | 27–29 | 26–28 | 26–29 | 21–31 | 24–28 | 25–28 | 29 | 29–31 | 26–29 | 24–26 | 24 | 24–28 |
Ventrolateral caudal tubercles anteriorly present (1) or not (0) | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 |
Subcaudal keeled (1) or smooth (0) | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 |
Single median row of keeled subcaudals (1) or smooth (0) | 1 | 1 | 0 | 0 | w | 0 | 1 | 0 | 0 | 0 | 1 | w |
Enlarge median subcaudal scales row (1) or not (0) | 0 | 0 | 1 | 0 | w | 0 | 0 | 1 | w | 1 | 0 | 1 |
Caudal tubercles restricted to a single paravertebral row |
1 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 0 |
No. of postcloacal tubercles in males | 1 | 1 | 1 | 1–2 | 1–2 | 1 | 2 | 1–3 | 1–2 | 1–2 | 0 | 1–3 |
Subtibial scales keeled (1) or smooth (0) | 1 | 1 | 1 | 0 | 0,1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Subcaudal region yellow present (1) or not (0) | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 0 |
Ventral pattern sexually dimorphic present (1) or not (0) | 1 | 1 | 1 | 1 | – | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Dorsal color pattern sexually dimorphic (1) or not (0) | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 |
Adult male;
Body slender, elongate (
Fore and hind limbs moderately long, slender; scales beneath forearm slightly raised, smooth and sub-imbricate; subtibial scales keeled; palmar scales smooth and juxtaposed; digits elongate, slender, inflected joint and bearing slightly recurved claws; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular; lamellae of distal phalanges wide; lamellae beneath inflection large; interdigital webbing absent; enlarged submetatarsal scales on 1st toe absent; total subdigital lamellae on fingers: 17-21-25-28-26 (right manus), 17–16 (broken)-25-28-26 (left manus); fingers increase in length from first to fourth with fourth and fifth nearly equal in length; relative length of fingers IV>V>III>II>I; total subdigital lamellae on toes: 13-21-24-29-25 (right pes), 13-21-24-29-25 (left pes); toes increase in length from first to fifth with fourth and fifth nearly equal in length; relative length of toes IV>V>III>II>I.
The original tail cylindrical, swollen at the base and longer than head and body (
(in mm; Table
Descriptive measurements in millimeters and characters of the type series of
Museum number | |||||||||||||||||||
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Type series | H | P | P | P | P | P | P | P | P | P | P | P | P | P | P | P | P | P | P |
Sex | M | M | M | M | M | M | M | M | M | M | M | M | F | F | F | F | F | F | F |
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40.1 | 39.5 | 40.1 | 37.8 | 37.6 | 39.1 | 37.7 | 36.7 | 37.0 | 39.4 | 40.6 | 39.9 | 37.3 | 38.3 | 38.8 | 41.8 | 41.4 | 37.5 | 41.1 |
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54.7 | 39.7r | 47.3r | 37.4r | 38.9r | 39.2r | 49.1 | 32.6r | 33.9r | 45.9r | 39.5r | 55.4 | 37.3r | 47.6r | 17.6b | 56.1 | 44.35r | 53.3 | 51.5 |
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3.9 | 4.0 | 4.1 | 3.8 | 3.7 | 4.0 | 3.6 | 3.7 | 3.8 | 3.9 | 4.1 | 3.9 | 3.6 | 3.8 | 3.6 | 4.2 | 4.0 | 3.8 | 3.9 |
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5.8 | 5.8 | 5.8 | 5.6 | 5.6 | 5.7 | 5.7 | 5.5 | 5.6 | 5.7 | 5.8 | 5.8 | 5.6 | 5.7 | 5.6 | 5.8 | 5.7 | 5.6 | 5.8 |
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7.2 | 7.2 | 7.3 | 7.2 | 7.1 | 7.2 | 7.2 | 7.1 | 7.1 | 7.2 | 7.3 | 7.2 | 7.1 | 7.1 | 7.2 | 7.3 | 7.3 | 7.1 | 7.2 |
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17.4 | 17.5 | 17.6 | 17.3 | 17.3 | 17.5 | 17.3 | 16.8 | 16.9 | 17.4 | 17.6 | 17.5 | 17.4 | 17.5 | 17.4 | 17.6 | 17.5 | 17.3 | 17.5 |
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10.3 | 10.2 | 10.3 | 10.0 | 10.1 | 10.2 | 10.1 | 10.1 | 10.2 | 10.3 | 10.4 | 10.4 | 9.9 | 10.2 | 10.1 | 10.4 | 10.4 | 10.0 | 10.4 |
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6.3 | 6.2 | 6.3 | 6.1 | 6.1 | 6.2 | 6.1 | 6.0 | 6.0 | 6.2 | 6.3 | 6.3 | 6.1 | 6.2 | 6.2 | 6.3 | 6.2 | 6.1 | 6.3 |
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4.2 | 4.2 | 4.2 | 4.0 | 3.8 | 4.2 | 3.8 | 3.9 | 4.0 | 4.1 | 4.2 | 4.1 | 3.8 | 4.0 | 3.9 | 4.3 | 4.2 | 4.0 | 4.2 |
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2.4 | 2.3 | 2.4 | 2.2 | 2.2 | 2.3 | 2.1 | 2.1 | 2.2 | 2.3 | 2.4 | 2.3 | 2.1 | 2.2 | 2.3 | 2.4 | 2.4 | 2.3 | 2.4 |
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3.1 | 3.0 | 3.1 | 2.8 | 2.8 | 3.0 | 2.9 | 2.8 | 2.9 | 3.1 | 3.1 | 3.1 | 2.9 | 3.0 | 2.9 | 3.2 | 3.2 | 3.1 | 3.2 |
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4.8 | 4.9 | 5.1 | 4.7 | 4.7 | 4.9 | 4.7 | 4.7 | 4.7 | 5.1 | 5.1 | 4.8 | 4.6 | 4.8 | 4.7 | 5.1 | 5.1 | 4.8 | 5.1 |
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3.9 | 3.6 | 3.8 | 3.4 | 3.4 | 3.5 | 3.4 | 3.6 | 3.6 | 3.8 | 3.9 | 3.8 | 3.4 | 3.7 | 3.7 | 3.9 | 3.9 | 3.7 | 3.8 |
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2.9 | 2.8 | 2.9 | 2.8 | 2.8 | 2.9 | 2.6 | 2.7 | 2.8 | 2.8 | 2.9 | 2.8 | 2.6 | 2.7 | 2.6 | 2.9 | 2.9 | 2.7 | 2.9 |
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1.0 | 1.1 | 1.1 | 1.0 | 1.0 | 1.0 | 1.0 | 0.9 | 0.9 | 1.1 | 1.1 | 1.0 | 1.0 | 1.0 | 0.9 | 1.0 | 1.0 | 1.0 | 1 |
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1.0 | 1.1 | 1.0 | 1.0 | 1.0 | 1.0 | 1.0 | 1.0 | 0.9 | 1.0 | 1.1 | 1.0 | 1.0 | 1.1 | 1.0 | 1.1 | 1.1 | 1.1 | 1.1 |
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9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 8 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 |
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9 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 9 | 8 | 8 | 9 | 9 | 9 | 9 | 9 | 9 | 9 |
Pore-bearing precloacal scales | 6 | 4 | 4 | 4 | 5 | 4 | 4 | 4 | 5 | 5 | 7 | 4 | – | – | – | – | – | – | – |
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19 | 20 | 20 | 19 | 20 | 20 | 20 | 19 | 20 | 21 | 20 | 20 | 19 | 21 | 20 | 19 | 20 | 21 | 19 |
Spine-like tubercles on flank | 5 | 5 | 6 | 5 | 6 | 6 | 6 | 6 | 6 | 5 | 6 | 6 | 6 | 6 | 5 | 5 | 6 | 6 | 5 |
4th toe lamellae | 29 | 29 | 29 | 28 | 28 | 28 | 27 | 28 | 28 | 27 | 27 | 29 | 29 | 27 | 27 | 28 | 27 | 28 | 27 |
(Fig.
Coloration of
Male paratypes of
Most paratypes approximate the holotype in general aspects of morphology (Figs
Female paratypes of
Habitats of
Seven specimens (
The male holotype was found during the night (1943 h) perched head down on a vertical surface in a crevice of a granitic rock boulder near a stream. A female paratype (
Paratypes that were found during the day were in shaded areas, crevices of boulders, rock walls and on boulder outcrops near streams. Paratypes found at night were in shaded surfaces of the boulders, within deep crevices, or perched on vegetation near a rocky stream. Three gravid females (
Ecological parameters of activity period, elevation (lowland < 600 m), microhabitat preference and presence or absence of ocelli (eyespots) in 18 species of
Species/ Parameters | Activity period | Elevation | Microhabitat preference | Ocelli location | |||||||
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Diurnal | Nocturnal | Lowland | Upland | Granite | Limestone | Vegetation | Terrestrial | Head | Neck | Shoulders | |
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The specific epithet
The complex geological history of Thailand created a large number of granitic rocky outcrop ecosystems in southern Thailand (Charusiri 1993; Cobbing et al. 2011). This ecosystem supports high levels of species endemism and species diversity of gekkonid lizards, especially species in the genus
A decade ago, only four species of
The new species is known only from the type locality and likely has a narrow geographic distribution. It is expected to be found in other nearby granitic rocky streams in Kam Lon Subdistrict, Lan Saka District, Nakhon Si Thammarat Province. However, additional surveys for this species are needed to clarify the geographic range of the new species. Our findings agree with those of
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-PG-160022). AA was supported by Department of Zoology and International SciKU Branding (ISB), Faculty of Science, Kasetsart University. This research was approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project number ACKU60-SCI-004). Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Attapol Rujirawan, Korkhwan Termprayoon, Siriporn Yodthong and Piyawan Puenprapai assisted with fieldwork. Evan S. H. Quah and Vinh Q. Luu improved the manuscript. This paper is contribution number 917 of the Auburn University Museum of Natural History.
List of comparative specimens examined.