Corresponding author: Anchalee Aowphol (fsciacl@ku.ac.th)
Academic editor: Thomas Ziegler
Two new insular rock geckos in the genus
Ampai N, Rujirawan A, Yodthong S, Termprayoon K, Stuart BL, Wood Jr PL, Aowphol A (2022) Hidden diversity of rock geckos within the
The rock gecko genus
Historically, the taxonomy and systematics of Thai
We conducted field surveys for
Fieldwork was conducted at five different localities in southern Thailand including (1) Pathio District, Chumphon Province in June 2017, (2) Kanchanadit District, Surat Thani Province in January 2019, (3) Nayong District, Trang Province in May 2016, December 2017, and July 2019, (4) Ko Samui, Surat Thani Province in September 2015, July 2018, and August 2020, and (5) Ko Similan, Mu Ko Similan National Park, Phang-nga Province in March 2018. Sampling was conducted by using visual encounter surveys both during the day (1000–1800 h) and at night (1900–2200 h). Ecological data (air temperature and relative humidity) were collected using a Kestrel 4000 Weather Meter. Habitat preferences (e.g., microhabitat, substrate type and habitat use) were also recorded. Geographical coordinates and elevation were taken using a Garmin GPSMAP 64s. At each locality, specimens were photographed and euthanized by cardiac injection of tricaine methane sulfonate (MS-222) solution (
Genomic DNA from 21 individuals of
Voucher information, including locality, collection numbers, GenBank accession numbers and reference for the specimens used in the phylogenetic analyses. Voucher abbreviations are as follows:
Species | Locality | Collection number | GenBank accession number | Reference |
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Cambodia, Kampot |
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Thailand (captive) | No number |
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Cambodia, Pursat Province, Phnom Aural |
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Myanmar, Ayeyarwady Division, Myaungmya District |
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Cambodia, Pursat Province, Phnom Aural |
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Myanmar, Tanintharyi Division, Kaw Thaung District |
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Myanmar, Mon State, Kyait Hti Yo Wildlife Sanctuary |
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Vietnam, Ca Mau Province, Con Dao Archipelago |
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Vietnam, Ca Mau Province, Hon Khoai Island |
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Vietnam, Kien Giang Province, Hon Dat Hill |
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Vietnam, Kien Giang Province, Hon Tre Island |
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Cambodia, Pursat Province, Phnom Dalai |
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Thailand, Prachuap Khiri Khan Province, Kui Buri District, Wat Khao Daeng |
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Cambodia, Pursat Province, O’Lakmeas |
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Vietnam, An Giang Province, Nui Cam Hill |
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Malaysia, Perlis, Kuala Perlis |
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Malaysia, Perlis, Gua Kelam |
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Malaysia, Perlis, Perlis State Park |
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Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi |
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Thailand, Trang Province, Thum Khao Ting |
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Thailand, Satun Province, Mueang Satun District, Ko Tarutao |
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Malaysia, Terengganu, Gunung Lawit |
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Malaysia, Kelantan, Gunung Reng |
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Malaysia, Penang, Pulau Pinang |
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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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Malaysia, Kedah, Pulau Langkawi, Gunung Raya |
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Malaysia, Perak, Bukit Larut |
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Malaysia, Perak, Belum-Temengor, Sungai Enam |
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Thailand, Satun Province, Mueang Satun District, Ko Adang |
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This study | |||
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Thailand, Satun Province, Mueang Satun District, Ko Rawi |
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This study | ||
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Thailand, Trang Province, Nayong District |
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This study | |
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Thailand, Kanchanaburi Province, Sai Yok National Park |
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Thailand, Surat Thani Province, Kanchanadit District, Tai Rom Yen National Park |
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This study | |
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Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai Pak Waterfall |
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Thailand, Satun Province, Phuphaphet Cave |
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Malaysia, Perlis, Perlis State Park |
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Thailand, Phang-nga Province, Mueang Phang-nga 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, Surat Thani Province, Ko Samui District, Ko Samui, Hin Lad Waterfall |
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This study | ||
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Thailand, Phang-nga Province, Tai Mueang District, Mu Ko Similan National Park, Ko Similan, Ao Nguang Chang |
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This study | ||
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Thailand, Ratchaburi Province, Suan Phueng District, Khao Laem Mountain | ZMMU R 16391 |
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AUP 00767 |
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Thailand, Chumpon Province, Pathio District |
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This study | |||
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Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill |
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Thailand, Ranong Province, Suk Saran District, Naka |
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Homologous sequences of 68
The
Coloration and pattern in life was determined by examination of digital images taken of living specimens of all possible age classes prior to preservation. Morphological and meristic data were taken by the first author on the left side of preserved specimens for symmetrical characters, when possible, using digital Mitutoyo CD-6” ASX Digimatic Calipers to the nearest 0.1 mm under a Nikon SMZ745 dissecting microscope. Three body-size classes were established by snout-vent length: small size (< 35 mm), medium size (35–40 mm) and large size (> 40 mm). Only adult individuals, as determined by the presence of secondary sexual characteristics such as pore-bearing precloacal scales or hemipenes in males or visible eggs on ventral side of body or enlarged endolymphatic glands in females, were included for morphometric and meristic measurements. A total of sixteen morphological characters was scored following
Meristic characters states of scales and quantitative observations of pattern and structures were evaluated under a Nikon SMZ745 dissecting microscope. Meristic characters taken were modified from
Statistical analyses were used to compare differences in size and shape within the
Xadj = log(
where Xadj is the corrected value of the morphometric variable;
Univariate analyses were implemented in the Paleontological statistics software (PAST v4.07b;
The aligned dataset contained 1,310 characters of 89 individuals of
The single best tree from 10,000 Maximum likelihood bootstrap replicates based on 1,310 bp of the mitochondrial NADH dehydrogenase subunit 2 (
Mean (min-max) uncorrected pairwise distances (%) within the
Species | N | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 |
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1. |
4 |
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2. |
4 | 11.50 |
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(11.32–11.56) |
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3. |
9 | 9.32 | 11.12 |
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(8.87–9.68) | (11.08–11.18) |
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4. |
4 | 8.92 | 9.40 | 7.80 |
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(8.86–9.10) | (9.34–9.58) | (7.52–8.15) |
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5. |
3 | 23.00 (22.34–23.67) | 23.19 (22.72–24.10) | 22.47 (22.01–23.27) | 22.63 (22.10–23.58) |
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6. |
5 | 9.55 (9.10–9.73) | 11.73 (11.72–11.80) | 9.08 (8.86–9.44) | 8.38 (8.23–8.54) | 23.48 (23.13–24.10) |
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7. |
4 | 14.61 (14.10–14.96) | 16.15 (15.92–16.39) | 14.19 (13.63–14.77) | 14.04 (13.63–14.41) | 24.13 (23.02–25.41) | 14.55 (14.23–14.89) |
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8. |
2 | 10.98 (10.17–11.81) | 11.42 (10.79–12.06) | 8.20 (6.77–9.40) | 9.36 (8.72–9.96) | 24.74 (24.00–25.60) | 10.81 (10.04–11.65) | 16.15 (15.07–17.04) |
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9. |
2 | 9.68 (9.58–9.81) | 8.16 (8.16–8.16) | 9.00 (8.87–9.21) | 7.48 (7.36–7.75) | 21.69 (21.37–22.29) | 10.23 (10.06–10.36) | 15.62 (15.24–16.01) | 10.50 (9.70–11.33) |
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10. |
2 | 23.53 (21.83–24.93) | 25.24 (23.67–26.82) | 23.19 (21.75–24.53) | 23.74 (22.46–24.93) | 16.70 (15.57–17.59) | 24.34 (23.02–25.62) | 23.72 (21.82–25.62) | 24.79 (22.75–27.82) | 24.21 (22.78–25.65) |
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11. |
3 | 10.57 (10.21–10.83) | 11.21 (10.61–11.53) | 7.69 (7.04–8.20) | 8.83 (8.31–9.36) | 24.15 (23.27–24.50) | 10.23 (9.64–10.50) | 14.94 (13.80–15.76) | 8.34 (7.13–8.99) | 9.00 (8.39–9.36) | 23.14 (21.83–24.03) |
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12. |
2 | 22.75 (22.00–23.15) | 22.69 (22.48–22.91) | 21.53 (21.13–21.88) | 21.32 (20.97–21.72) | 16.08 (15.44–17.00) | 22.06 (21.84–22.28) | 22.46 (21.91–22.99) | 23.33 (23.02–23.57) | 21.35 (21.05–21.66) | 15.93 (15.29–17.13) | 22.15 (21.21–22.89) |
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13. |
4 | 22.04 (20.83–23.59) | 23.21 (21.73–24.88) | 22.20 (21.21–23.39) | 21.70 (20.64–23.11) | 19.28 (18.73–19.59) | 22.27 (21.12–23.83) | 23.00 (22.30–24.64) | 23.30 (21.21–24.58) | 21.68 (20.25–23.29) | 19.49 (18.12–20.70) | 23.26 (21.41–24.87) | 19.10 (18.72–19.55) |
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14. |
3 | 23.10 (22.66–23.29) | 24.27 (24.12–24.50) | 23.43 (22.97–23.89) | 22.59 (22.35–22.89) | 20.98 (20.79–21.11) | 23.04 (22.91–23.19) | 24.08 (23.65–24.58) | 25.03 (24.88–25.24) | 23.44 (23.29–23.71) | 20.83 (20.05–21.57) | 24.79 (24.09–25.32) | 20.73 (20.49–21.00) | 14.31 (13.86–15.21) |
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15. |
1 | 26.76 (26.56–26.83) | 27.11 (27.11–27.11) | 26.36 (26.15–26.42) | 26.49 (26.42–26.56) | 20.75 (20.66–20.80) | 27.52 (27.52–27.52 | 25.23 (24.65–25.86) | 27.36 (27.11–27.60) | 26.71 (26.59–26.83) | 20.89 (20.89–20.89) | 27.71 (27.66–27.80) | 23.24 (23.00–23.48) | 13.84 (13.64–14.21) | 17.45 (17.22–17.63) |
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Univariate analysis of variance (
Pairwise significant difference matrix from 15 size-corrected morphometric measurements of
No. | Species | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 |
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1 | – | ||||||||
2 | – | ||||||||
3 |
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4 |
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5 |
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6 |
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7 |
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8 |
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9 |
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Summary of proportions of variance, standard deviation, eigenvalues and factor loadings from the 10 first principal components (PC) of 14 size-adjusted morphometric characters of two new insular species
Character |
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PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 |
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Proportion of Variance | 54.6 | 17.2 | 6.2 | 5.7 | 5.1 | 4.3 | 2.3 | 1.3 | 1.0 | 0.8 |
Standard deviation | 2.77 | 1.55 | 0.93 | 0.89 | 0.84 | 0.78 | 0.57 | 0.42 | 0.37 | 0.33 |
eigenvalues | 7.65 | 2.406 | 0.872 | 0.793 | 0.711 | 0.606 | 0.32 | 0.179 | 0.137 | 0.112 |
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-0.293 | -0.083 | 0.266 | 0.296 | -0.093 | 0.017 | - |
0.069 | -0.054 | 0.169 |
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-0.263 | -0.125 | - |
- |
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-0.207 | - |
-0.01 | 0.005 |
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- |
0.091 | -0.217 | -0.171 |
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0.182 | 0.021 | 0.123 | 0.217 | - |
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0.233 | -0.04 | -0.132 | -0.093 | 0.247 | 0.069 | - |
- |
0.163 |
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-0.118 | -0.063 | -0.020 | 0.037 |
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0.229 |
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0.064 |
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- |
-0.097 | -0.009 | -0.030 | -0.115 |
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0.100 | 0.016 | -0.145 | -0.267 |
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- |
-0.073 |
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0.211 | -0.114 | 0.066 | 0.175 | 0.170 | -0.108 | 0.095 |
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-0.264 | 0.079 |
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-0.232 | 0.237 | - |
0.161 | 0.132 | - |
-0.196 |
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- |
-0.005 | -0.032 |
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0.145 | -0.073 | 0.080 | 0.071 | 0.209 | -0.164 |
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- |
0.191 | 0.011 | 0.198 | -0.086 | - |
0.098 | - |
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-0.027 |
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-0.210 | 0.208 | - |
0.093 | - |
- |
0.119 |
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-0.236 | 0.083 |
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-0.105 |
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0.282 | - |
- |
0.087 | -0.158 | 0.214 |
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0.004 |
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-0.096 | - |
0.172 | -0.231 | -0.137 | -0.155 |
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-0.085 | 0.254 | 0.292 |
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0.113 |
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0.14 | 0.211 |
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0.079 | 0.236 | 0.131 | 0.032 |
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Multivariate analysis results of principal component analysis (
The Ko Samui and Ko Similan populations distinctly differed from all congeners in the
Adult male holotype of
Coloration of
An adult male in good state of preservation; 42.3 mm
Body relatively slender, elongate (
Fore and hind limbs moderately elongate, slender; scales beneath forearm slightly raised, smooth and subimbricate; subtibial scales keeled; palmar scales smooth, flat and subimbricate; digits long, slender, distinctly inflected joint with strong, 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 present; total subdigital lamellae on fingers I–V: 18-21-22-24-23 (right manus), 18-21-22-24-23 (left manus); fingers increase in length from first to fourth with fifth nearly equal in length as fourth; relative length of fingers IV>V>III>II>I; total subdigital lamellae on toes I–V: 14-20-21-24-23 (right pes), 14-(broken)-21-24-23 (left pes); toes increase in length from first to fourth with fifth nearly equal in length as fourth; relative length of toes IV>V>III>II>I.
Tail complete, entire cylindrical, relatively slender, swollen at the base; tail length (
(in mm; Table
Descriptive measurements in millimeters and characters of the type series of
Characters/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 |
Sex | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Male | Female | Female | Female |
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42.3 | 40.1 | 41.2 | 40.8 | 37.0 | 38.7 | 41.7 | 41.5 | 40.7 | 41.0 | 40.4 | 35.6 | 40.1 | 36.7 | 40.6 | 36.4 | 41.2 | 41.6 |
Tail | C | C | C | R | B | C | R | C | R | B | B | C | C | C | C | C | R | R |
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52.2 | 50.8 | 57.8 | 56.2 | 44.3 | 48.8 | 51.3 | 59.3 | 40.6 | 28.7 | – | 47.6 | 54.8 | 48.6 | 59.6 | 46.1 | 44.0 | 16.4 |
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4.4 | 3.9 | 3.8 | 4.0 | 3.9 | 4.0 | 4.2 | 4.2 | 4.1 | 4.3 | 4.4 | 3.6 | 4.1 | 3.7 | 4.1 | 3.6 | 4.2 | 3.9 |
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6.5 | 6.3 | 6.3 | 6.2 | 6.2 | 6.1 | 6.3 | 6.2 | 6.1 | 6.5 | 6.8 | 6.2 | 6.2 | 6.0 | 6.1 | 6.2 | 6.2 | 6.1 |
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7.9 | 7.5 | 7.6 | 7.6 | 7.5 | 7.5 | 7.7 | 7.6 | 7.7 | 7.6 | 7.8 | 7.2 | 7.6 | 6.9 | 7.7 | 7.1 | 7.8 | 7.7 |
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17.9 | 17.7 | 17.7 | 17.7 | 16.1 | 16.1 | 17.7 | 17.6 | 17.7 | 17.7 | 17.6 | 15.5 | 17.6 | 16.1 | 17.8 | 15.2 | 17.9 | 17.9 |
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11.5 | 11.2 | 11.3 | 11.2 | 10.8 | 10.8 | 11.4 | 11.6 | 10.7 | 11.4 | 11.1 | 9.8 | 10.8 | 10.0 | 11.4 | 10.6 | 11.3 | 10.8 |
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6.9 | 6.6 | 6.9 | 6.8 | 6.6 | 6.6 | 6.8 | 6.8 | 6.8 | 6.8 | 6.7 | 6.2 | 6.7 | 6.2 | 6.9 | 6.2 | 6.9 | 6.8 |
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4.5 | 4.2 | 4.3 | 4.2 | 4.1 | 4.1 | 4.4 | 4.3 | 4.3 | 4.4 | 4.4 | 3.8 | 4.3 | 3.9 | 4.4 | 3.8 | 4.4 | 4.4 |
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2.5 | 2.3 | 2.4 | 2.2 | 2.2 | 2.3 | 2.4 | 2.4 | 2.1 | 2.2 | 2.2 | 2.3 | 2.2 | 2.2 | 2.4 | 2.2 | 2.4 | 2.4 |
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3.3 | 3.1 | 3.3 | 3.2 | 3.1 | 3.1 | 3.2 | 3.2 | 3.1 | 3.2 | 3.2 | 2.9 | 3.2 | 3.0 | 3.2 | 2.9 | 3.2 | 3.2 |
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5.0 | 4.9 | 4.9 | 4.9 | 4.5 | 4.6 | 4.8 | 4.8 | 4.7 | 4.9 | 4.9 | 4.6 | 4.9 | 4.7 | 4.8 | 4.5 | 4.9 | 4.9 |
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4.0 | 3.7 | 3.7 | 3.7 | 3.8 | 3.7 | 4.0 | 3.9 | 3.7 | 3.8 | 3.7 | 3.4 | 3.6 | 3.5 | 3.9 | 3.5 | 4.0 | 3.9 |
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1.0 | 0.9 | 0.8 | 0.8 | 0.9 | 0.7 | 0.9 | 1.0 | 1.0 | 0.9 | 0.9 | 0.9 | 0.8 | 0.9 | 0.9 | 0.9 | 0.9 | 0.8 |
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3.3 | 3.1 | 3.3 | 3.2 | 3.0 | 3.2 | 3.2 | 3.1 | 3.1 | 3.2 | 3.1 | 2.8 | 3.1 | 2.8 | 3.2 | 2.8 | 3.3 | 3.3 |
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1.1 | 1.0 | 1.1 | 1.0 | 0.9 | 1.0 | 1.1 | 1.0 | 1.0 | 1.0 | 1.0 | 1.0 | 0.9 | 1.0 | 1.1 | 1.0 | 1.0 | 1.1 |
Supralabial scales | 9 | 9 | 9 | 9 | 9 | 9 | 8 | 9 | 8 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 |
Infralabial scales | 8 | 9 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 9 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
No. of precloacal pores | 8 | 7 | 8 | 5 | 6 | 8 | 7 | 8 | 7 | 6 | 7 | 8 | 7 | 5 | 7 | – | – | – |
Precloacal pore continuous (1) or separated (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | – | – | – |
Precloacal pores elongate (1) or round (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | – | – | – |
No. of paravertebral tubercles | 26 | 26 | 25 | 27 | 26 | 27 | 27 | 26 | 27 | 25 | 27 | 26 | 25 | 27 | 27 | 27 | 27 | 26 |
Tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
Tubercles present (1) or absent (0) on lower flanks | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
No. of 4th toe lamellae | 24 | 23 | 25 | 25 | 25 | 25 | 23 | 25 | 24 | 24 | 25 | 22 | 25 | 25 | 25 | 24 | 24 | 25 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Pectoral scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Ventral scales on thigh keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Subcaudal keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Caudal tubercles restricted to the single paravertebral row on each side (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Adult male holotype of
Paratypes of
Most paratypes closely resemble the holotype in all aspect of pattern and coloration. Morphometric and meristic variation within the type series is presented in Table
Habitats of
The type locality is surrounded by lowland evergreen forest with granitic rocky outcrops along Lipa Yai Canal in the western part of Ko Samui. All specimens of
The specific epithet
Meristic character state and color pattern of species in the
Characters / Species |
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|
||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Sample size | 18 | 4 | 15 | 25 | 5 | 4 | 19 | 8 | 2 | 5 | 2 | 12 | 6 | 3 |
Maximum |
42.3 | 48.1 | 44.9 | 40.9 | 43.5 | 37.8 | 41.8 | 41.3 | 42.0 | 49.6 | 36.2 | 39.7 | 39.0 | 44.7 |
Supralabial scales | 8 or 9 | 8 or 9 | 10 | 8–10 | 7–10 | 8 or 9 | 9 | 8 or 9 | 10 | 8 | 10 or 11 | 8 or 9 | 10 or 11 | 8 or 9 |
Infralabial scales | 8 or 9 | 7 or 8 | 9 | 8 | 6–9 | 7 or 8 | 9 | 7 or 8 | 10 | 7 or 8 | 10 | 6–8 | 9–11 | 7–9 |
No. of pore-bearing precloacal scales | 5–8 | 1 | 6–8 | 6–8 | 5–8 | 6 or 7 | 4–7 | 3–6 | 4 | 0 | 6 or 7 | 0 | 0 | 4 |
Pore-bearing precloacal scales row continuous (1) or separated (0) | 0 | – | 0 | 0 | 1 | 1 | 0 | 0 | 1 | – | 1 | – | – | 0 |
Pore-bearing precloacal scales elongate (1) or round (0) shapes | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | – | 1 | – | – | 0 |
No. of paravertebral tubercles | 25–27 | 24 or 25 | 23–25 | 22–25 | 18–24 | 19–24 | 19–21 | 22–29 | 22 | 24–27 | 16–18 | 19–25 | 15–19 | 25–29 |
Paravertebral tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 | w or 0 | w | 1 | w or 0 | 1 | w | 0 | 0 | 1 | 0 |
Tubercles present (1) or absent (0) on lower flanks | 1 | 1 | 0 | 1 | 1 | 1 | 1 | w or 1 | 0 | 1 | 0 | 1 | 1 | 1 |
No. of 4th toe lamellae | 22–25 | 23 or 24 | 26–28 | 26–29 | 21–31 | 24–28 | 27–29 | 25–28 | 29 | 29–31 | 22 | 24–26 | 24 | 24–28 |
Ventral scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 0 | w or 0 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 1 |
Subcaudal scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 1 | 1 | 1 |
Single median row of keeled subcaudals (1) or smooth (0) scales | 1 | 1 | 1 | 0 | 0 | w | 1 | 0 | 1 | 0 | – | 0 | 1 | w |
Enlarged median subcaudal scales row (1) or not (0) | 1 | 0 | 0 | 1 | 0 | w | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 1 |
Caudal tubercles restricted to a single paravertebral row on each side (1) or not (0) | 0 | 0 | 0 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | – | 0 | 1 | 0 |
Ventrolateral caudal tubercles anteriorly present (1) or not (0) | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 |
No. of postcloacal tubercles in males | 1 or 2 | 2 | 1 | 1 | 1 or 2 | 1 or 2 | 1 | 1 | 2 | 1–3 | 2 | 1 or 2 | 0 | 1–3 |
Subtibial scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 | 0 | 0 or 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 |
Yellow coloration in the subcaudal region present (1) or not (0) | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
Ventral pattern sexually dimorphic present (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | – | 1 | 1 | 1 |
Adult male holotype of
Coloration of adult
An adult male in good state of preservation; 47.6 mm
Body robust, not elongate (
Fore and hind limbs moderately long, slender; scales beneath forearm slightly raised, smooth and subimbricate; subtibial scales keeled; palmar scales keeled, flat and subimbricate; digits long, slender with inflected joint; claws slightly recurved; 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 generally absent; enlarged submetatarsal scales on 1st toe present; total subdigital lamellae on fingers I–V: 15-21-22-24-23 (right manus), 15-21-23-24-23 (left manus); fingers increase in length from first to fourth with fifth nearly equal in length as fourth; relative length of fingers IV>V>III>II>I; total subdigital lamellae on toes I–V: 17-20-22-24-23 (right pes), 17-19-22-24-23 (left pes); toes increase in length from first to fourth with fifth nearly equal in length as fourth; relative length of toes IV>V>III>II>I.
Tail regenerated, subcylindrical, relatively swollen at the base; tail length (
(in mm; Table
Descriptive measurements in millimeters and characters of the type series of
Characters / Museum number | ||||
---|---|---|---|---|
Sex | Male | Female | Female | Female |
Type series | H | P | P | P |
|
47.6 | 48.1 | 38.6 | 44.2 |
Tail | R | R | R | C |
|
49.6 | 43.2 | 37.6 | 58.2 |
|
4.6 | 4.6 | 4.1 | 4.4 |
|
6.8 | 6.9 | 6.2 | 6.6 |
|
8.6 | 8.8 | 7.5 | 8.4 |
|
19.6 | 19.8 | 16.6 | 19.4 |
|
12.4 | 12.6 | 10.4 | 12.1 |
|
7.8 | 7.9 | 6.5 | 7.7 |
|
4.8 | 4.9 | 4.1 | 4.6 |
|
2.4 | 2.4 | 2.1 | 2.3 |
|
3.6 | 3.7 | 3.0 | 3.4 |
|
5.3 | 5.4 | 4.3 | 4.9 |
|
4.0 | 4.1 | 3.4 | 3.7 |
|
1.0 | 1.0 | 0.9 | 1.0 |
|
3.1 | 3.2 | 2.6 | 2.6 |
|
1.1 | 1.1 | 0.9 | 1.0 |
Supralabial scales | 8 | 9 | 9 | 9 |
Infralabial scales | 7 | 8 | 8 | 8 |
No. of precloacal pores | 1 | – | – | – |
Precloacal pore continuous (1) or separated (0) | – | – | – | – |
Precloacal pores elongate (1) or round (0) | 0 | – | – | – |
No. of paravertebral tubercles | 24 | 25 | 25 | 24 |
Tubercles linearly arranged (1) or more random (0) | 0 | 0 | 0 | 0 |
Tubercles present (1) or absent (0) on lower flanks | 1 | 1 | 1 | 1 |
No. of 4th toe lamellae | 24 | 24 | 23 | 23 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | 1 |
Pectoral scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 |
Ventral scales on thigh keeled (1) or smooth (0) | 1 | 1 | 1 | 1 |
Subcaudal keeled (1) or smooth (0) | 1 | 1 | 1 | 1 |
Subtibial scales keeled (1) or smooth (0) | 1 | 1 | 1 | 1 |
Enlarged median subcaudal scale row (1) or not (0) | 0 | 0 | 0 | 0 |
Caudal tubercles restricted to the single paravertebral row on each side (1) or not (0) | 1 | 1 | 1 | 1 |
Adult male holotype of
Due to having only a single adult male (
Habitats of
The type locality is dominated by mixed evergreen forest with shrub and beach forests. Ao Nguang Chang Bay is located at the southern part of the largest island, Ko Similan (= Ko Pad). All specimens of
The specific epithet
Historically, most Thai
This study revealed two unrecognized species of
This work was financially supported by Srinakharinwirot University Research Grant (No. 596/2564) and the Thailand Research Fund (DBG6080010). AR and AA were supported by Kasetsart University Research and Development Institute (KURDI), Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation (No. RGNS 64-038) and the Department of Zoology, Faculty of Science, Kasetsart University. Department of National Parks, Wildlife and Plant Conservation, Thailand provided permission to conduct the research (Permit no.0907.4/20800 and 0907.4/28401). This research was approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project no. ACKU61-SCI-008). We thank Ruamsilp Manajongprasert (Mu Ko Similan National Park) for facilitating the fieldwork. Piyawan Puanprapai and Akrachai Aksornneam assisted with fieldwork. Lee Grismer and an anonymous reviewer improved the manuscript.
Table S1
excel file
Significant