Corresponding author: Attapol Rujirawan (fsciapr@ku.ac.th)
Academic editor: Thomas Ziegler
Species delimitation and species diagnosis must remain separate operations to avoid constructing taxonomies comprised of non-monophyletic species based on morphological similarity as opposed to phylogenetic propinquity. This is particularly true for highly specialized species such as the range-restricted upland taxa in the
Grismer LL, Aowphol A, Yodthong S, Ampai N, Termprayoon K, Aksornneam A, Rujirawan A (2022) Integrative taxonomy delimits and diagnoses cryptic arboreal species of the
A cornerstone of biodiversity conservation is a phylogenetic-based taxonomy where the names of the component species are consistent with the patterns and processes by which they evolved. Taxonomies constructed from paraphyletic or polyphyletic species misrepresent history, thus obscuring true diversity and potentially countermanding the effectiveness of conservation efforts. Before a newly discovered population can be given a new name, or the name of an existing species be successfully challenged, it should be properly delimited and diagnosed. Delimiting and diagnosing species are independent operations used together to construct taxonomies that reflect, and are consistent with evolutionary history. Unfortunately, these two operations are often conflated when analyses to diagnose species, which are most often rooted in morphological similarity, are equated with analyses to delimit species, which are rooted in phylogenetic propinquity. The unfortunate consequence of this is that taxonomies may be constructed using non-monophyletic species, thus obscuring rather than revealing the group’s actual diversity. This is especially true for taxonomies comprised of highly specialized cryptic species where, in the absence of a phylogeny, morphological convergence can be mistaken for common ancestry (see
Convergent morphology results from the specific resource requirements that necessitate a particular functional morphology (
Distribution of nominal species and unnamed populations and specimens of the
The phylogenetic-based taxonomy of
The general lineage concept (
Species boundaries were cross-checked using a Bayesian Poisson Tree Process for species delimitation (
Methods for DNA extraction, sequencing, and editing followed
Species | Location | Catalog no. | GenBank no. |
---|---|---|---|
|
Thailand, no data | LSUHC 1899 | not in tree |
|
Thailand, Nakhon Si Thammarat Province, Nopphitam District, Khao Nan National Park, Huay Lak Protected Unit | THNHM 10670 | not in tree |
|
Thailand, Nakhon Si Thammarat Province, Lan Saka District, Khao Luang National Park | THNHM 14112 | not in tree |
|
Thailand, Nakhon Si Thammarat Province, Khao Ram Mt. | AUP-00573 |
|
|
Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | LSUHC 11788 | not in tree |
|
Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | LSUHC 15076 | not in tree |
|
Peninsular Malaysia, Pahang State, Fraser’s Hill, the Gap | ZRC 2.6091/LSUHC 6471 |
|
|
Peninsular Malaysia, Negeri Sembilan State | LSUHC 8238 | not in tree |
|
Peninsular Malaysia, Pahang State, near Cameron Highlands | LSUHC 12180 | not in tree |
|
Peninsular Malaysia, Pahang State, near Cameron Highlands | LSUHC 12181 | not in tree |
|
Malaysian pet trade, no data | ZMMU R-16144 | not in tree |
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00958 paratype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00959 holotype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00960 paratype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00961 paratype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00962 paratype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00963 paratype |
|
|
Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00964 paratype |
|
|
|
Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20226 paratype | not in tree |
|
Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20227 paratype | not in tree |
|
Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20228 paratype | not in tree |
|
Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20229 paratype | not in tree |
|
Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | YC000952 |
|
Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko Chang | ZMKU R 00945 holotype |
|
|
Thailand, Ranong Province, Khlong Nakha Wildlife Sanctuary | THNHM 01667 | not in tree | |
|
Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | HNUE-R00111 holotype |
|
|
Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | HNUE-R00112 paratype | not in tree |
|
Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | IEBR 4829 paratype |
|
|
Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | VNUF R.2020.12 paratype |
|
Laos, Khammouane Province | VNUF R.2014.50 |
|
|
Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 255454 |
|
|
Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 270492 |
|
|
Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 270493 | not in tree | |
Thailand, Loei Province, Nam San Noi River, Phu Luang Wildlife Sanctuary | FMNH 265806 |
|
|
Laos, Xaignabouli Province, Ban Pha Liep, Houay Liep Stream | NCSM 79472 |
|
|
Laos, Vientiane Province | ZMMU R-14917 | not in tree | |
Laos, Vientiane Province, tributary of Nam Pha River, Houay Wan Stream | NCSM 80100 |
|
|
Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U Waterfall | ZMKU R 00946 paratype |
|
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Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U Waterfall | ZMKU R 00947 holotype |
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|
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Thailand, Ratchaburi Province, Suan Phueng District, Khao Laem Mountain | ZMMU R-16851 holotype |
|
|
Thailand, Ratchaburi Province, Suan Phueng District, Hoop Phai Tong | ZMMU R-16852 paratype | not in tree |
|
Thailand, Ratchaburi Province, Suan Phueng District, Suan Phueng Subdistrict | ZMKU R 00948 |
|
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 01807 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03251 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03252 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03253 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03254 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 24622 | not in tree |
|
Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 24838 | not in tree |
Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam Subdistrict | ZMKU R 00949 holotype |
|
|
Thailand, Kanchanaburi Province, Thong Pha Phum District, Thong Pha Phum National Park | AUP-01715 |
|
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Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00950 |
|
|
Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00951 |
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Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00952 |
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Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00953 |
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Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00954 |
|
|
Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00955 |
|
|
Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00956 |
|
|
Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00957 |
|
|
Thailand, Tak Province, Tha Song Yang District, Mae Moei National Park, Chao Doi Waterfall | AUP-00680 |
|
|
Thailand, Phitsanulok Province, Phu Hin Rong Kla National Park | ZMMU R-16492 |
|
|
Thailand, Tak Province, Umphang District, Thung Yai Naresuan Wildlife Sanctuary | THNHM 00104 | not in tree | |
Thailand, Kanchanaburi Province, Thong Pha Phum District, Ban Saphan Lao | THNHM 27821 | not in tree | |
Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | USMHC 2555 |
|
Morphological data included both meristic and morphometric characters. To reduce the degree of researcher bias, data were taken using the protocol of
Meristic characters evaluated were the number of
supralabial scales
An input file implemented in
All statistical analyses were conducted using
Small sample sizes (
Morphospatial clustering and positioning among the species/populations was analyzed using multiple factor analysis (
A non-parametric permutation multivariate analysis of variance (
The
Some of the populations examined had only genetic or morphological data. However, only phylogenetically delimited populations bearing morphological differences from other populations were described as new species. In some cases, populations represented by only morphological data but in close geographic proximity to named species from which they could not be separated morphologically, were considered
The
Maximum clade credibility
Uncorrected pairwise sequence divergences within the
Mean (minimum–maximum) percentages of uncorrected pairwise sequence divergence (
|
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21.03 |
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21.68 | 4.39 |
|
|||||||||||||
20.77 | 22.58 | 21.42 |
|
||||||||||||
18.86 | 10.64 | 11.02 | 20.15 |
|
|||||||||||
(18.84–18.97) | (10.58–10.84) | (10.97–11.23) | (20.13–20.26) |
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20.77 | 6.97 | 9.16 | 22.84 | 12.02 |
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(12.00–12.13) | |||||||||||||||
19.35 | 14.58 | 14.71 | 20.90 | 12.31 | 15.23 |
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(12.26–12.31) | |||||||||||||||
20.70 | 3.30 | 3.71 | 21.11 | 11.34 | 8.13 | 14.58 |
|
||||||||
(20.65–20.90) | (2.84–4.00) | (3.35–4.26) | (20.90–21.42) | (11.10–11.87) | (7.74–8.65) | (14.45–14.84) |
|
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20.00 | 15.87 | 15.03 | 21.61 | 12.57 | 15.48 | 12.26 | 15.03 |
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|||||||
(19.74–20.26) | (15.61–16.13) | (14.84–15.23) | (21.42–21.81) | (12.26–13.03) | (15.23–15.74) | (12.00–12.52) | (14.71–15.48) | ||||||||
20.65 | 15.42 | 15.48 | 21.61 | 12.25 | 16.00 | 13.10 | 15.23 | 4.65 |
|
||||||
(20.13–21.16) | (14.84–16.00) | (14.84–16.13) | (21.16–22.06) | (11.61–13.03) | (15.35–16.65) | (12.52–13.68) | (14.19–16.23) | (3.61–5.68) | |||||||
20.34 | 7.93 | 9.51 | 22.02 | 9.75 | 8.96 | 13.22 | 8.81 | 13.12 | 13.25 |
|
|||||
(20.13–20.65) | (7.74–8.00) | (9.42–9.55) | (21.81–22.32) | (9.55–9.94) | (8.77–9.03) | (13.03–13.29) | (8.13–9.68) | (12.77–13.42) | (12.52–13.94) |
|
|||||
19.87 | 9.29 | 10.84 | 21.94 | 10.12 | 10.19 | 13.68 | 10.21 | 13.94 | 14.32 | 8.06 |
|
||||
(10.06–10.32) | (10.06–10.45) | (13.68–14.19) | (13.68–14.97) | (7.87–8.13) | |||||||||||
20.39 | 7.23 | 8.90 | 22.19 | 11.12 | 3.87 | 14.58 | 8.28 | 15.35 1) | 15.61 | 8.96 | 10.45 |
|
|||
(11.10–11.23) | (8.00–8.65) | (15.10–15.6 | (14.97–16.26) | (8.77–9.03) | |||||||||||
6.45 | 20.90 | 20.65 | 20.00 | 18.34 | 20.13 | 19.10 | 20.52 | 19.74 | 20.00 | 19.60 | 18.84 | 19.61 |
|
||
(18.32–18.45) | (20.26–20.65) | (19.48–20.00) | (19.48–20.52) | (19.48–19.87) | |||||||||||
19.74 | 5.81 | 8.13 | 21.16 | 10.12 | 7.1 | 13.94 | 6.97 | 13.94 | 13.94 | 7.80 | 8.39 | 6.58 | 19.48 |
|
|
(10.06–10.32) | (6.58–7.61) | (13.68–14.19) | (13.29–14.58) | (7.61–7.87) |
The
Species delimited by the
Species |
|
---|---|
|
1 |
1 | |
0.967 | |
|
1 |
0.957 | |
|
0.997 |
1 | |
|
0.841 |
0.575 | |
0.947 | |
0.947 | |
1 | |
0.565 | |
1 | |
0.997 | |
1 |
The
The phylomorphospace analysis illustrates that the morphological similarities among the species/populations in the
Given the phylogenetic delimitation of the Si Sawat, Prachuap Khiri Khan, Uthai Thani, and Ko Chang populations (Figs
Adult male ZMKU R 00959 from Tham Than Lot Noi-Tham Than Lot Yai Nature Trail, Chaloem Rattanakosin National Park, Khao Chot Subdistrict, Si Sawat District, Kanchanaburi Province, Thailand (
Adult males ZMKU R 00958 and ZMKU R 00960 and adult females ZMKU R 00961–64 bear the same collection site as the holotype.
Holotype of
Body relatively short (
Forelimbs moderate in stature, relatively short (
Tail original, 97.6 mm long (
Type series of
The specific epithet
All individuals were found in karst forests bearing mixed deciduous and dry evergreen trees amidst rocky streams and a nearby waterfall (Fig.
Habitat of
Summary statistics of the normalized morphometric data for the putative species of the
Species |
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Mean | 1.82 | 1.49 | 0.96 | 0.96 | 1.06 | 1.02 | 1.27 | 1.10 | 0.87 | 0.67 | 0.75 | 0.86 | 0.73 | 0.68 | 0.07 | 0.32 |
SD | 0.020 | 0.027 | 0.031 | 0.023 | 0.024 | 0.036 | 0.012 | 0.006 | 0.017 | 0.041 | 0.025 | 0.008 | 0.019 | 0.043 | 0.058 | 0.041 |
Lower | 1.80 | 1.46 | 0.92 | 0.92 | 1.02 | 0.97 | 1.25 | 1.10 | 0.86 | 0.63 | 0.71 | 0.85 | 0.70 | 0.62 | -0.02 | 0.25 |
Upper | 1.85 | 1.52 | 0.99 | 0.98 | 1.09 | 1.07 | 1.29 | 1.11 | 0.89 | 0.73 | 0.77 | 0.87 | 0.75 | 0.73 | 0.13 | 0.36 |
Mean | 1.91 | 1.59 | 0.80 | 1.05 | 1.14 | 1.13 | 1.34 | 1.20 | 1.00 | 0.71 | 0.83 | 0.95 | 0.79 | 0.70 | 0.19 | 0.43 |
SD | 0.016 | 0.012 | 0.341 | 0.026 | 0.022 | 0.024 | 0.001 | 0.012 | 0.008 | 0.017 | 0.020 | 0.008 | 0.010 | 0.070 | 0.062 | 0.033 |
Lower | 1.89 | 1.57 | 0.34 | 1.02 | 1.11 | 1.10 | 1.34 | 1.18 | 0.98 | 0.68 | 0.81 | 0.94 | 0.78 | 0.61 | 0.15 | 0.40 |
Upper | 1.93 | 1.60 | 1.09 | 1.08 | 1.16 | 1.15 | 1.34 | 1.21 | 1.00 | 0.72 | 0.85 | 0.95 | 0.80 | 0.76 | 0.28 | 0.48 |
Mean | 1.86 | 1.52 | 0.95 | 1.01 | 1.11 | 1.03 | 1.31 | 1.14 | 0.91 | 0.69 | 0.78 | 0.92 | 0.79 | 0.73 | 0.20 | 0.38 |
SD | 0.027 | 0.009 | 0.013 | 0.017 | 0.018 | 0.012 | 0.007 | 0.011 | 0.011 | 0.007 | 0.016 | 0.008 | 0.011 | 0.006 | 0.04 | 0.02 |
Lower | 1.82 | 1.51 | 0.94 | 0.98 | 1.10 | 1.01 | 1.30 | 1.13 | 0.90 | 0.68 | 0.76 | 0.91 | 0.78 | 0.72 | 0.14 | 0.35 |
Upper | 1.89 | 1.54 | 0.98 | 1.04 | 1.15 | 1.05 | 1.32 | 1.16 | 0.93 | 0.70 | 0.81 | 0.94 | 0.81 | 0.74 | 0.24 | 0.40 |
Mean | 1.86 | 1.50 | 0.98 | 1.01 | 1.10 | 1.08 | 1.30 | 1.12 | 0.80 | 0.72 | 0.76 | 0.90 | 0.77 | 0.68 | 0.13 | 0.33 |
SD | 0.060 | 0.016 | 0.032 | 0.015 | 0.020 | 0.007 | 0.012 | 0.006 | 0.187 | 0.026 | 0.026 | 0.024 | 0.034 | 0.0307 | 0.033 | 0.035 |
Lower | 1.78 | 1.48 | 0.94 | 0.99 | 1.07 | 1.07 | 1.28 | 1.12 | 0.56 | 0.69 | 0.73 | 0.88 | 0.74 | 0.66 | 0.10 | 0.29 |
Upper | 1.91 | 1.52 | 1.01 | 1.02 | 1.12 | 1.09 | 1.31 | 1.13 | 0.96 | 0.75 | 0.79 | 0.94 | 0.81 | 0.73 | 0.17 | 0.38 |
Value | 1.78 | 1.51 | 1.00 | 0.95 | 1.05 | 1.02 | 1.30 | 1.14 | 0.91 | 0.69 | 0.76 | 0.91 | 0.78 | 0.68 | 0.05 | 0.36 |
Value | 1.85 | 1.50 | 1.02 | 0.94 | 1.08 | 1.08 | 1.27 | 1.09 | 0.90 | 0.72 | 0.70 | 0.88 | 0.75 | 0.60 | 0.12 | 0.36 |
Mean | 1.83 | 1.47 | 0.91 | 0.99 | 1.06 | 1.05 | 1.31 | 1.08 | 0.85 | 0.57 | 0.76 | 0.86 | 0.81 | 0.74 | -0.12 | 0.43 |
SD | 0.009 | 0.002 | 0.004 | 0.007 | 0.000 | 0.006 | 0.001 | 0.003 | 0.010 | 0.035 | 0.016 | 0.015 | 0.004 | 0.009 | 0.019 | 0.008 |
Lower | 1.82 | 1.47 | 0.91 | 0.98 | 1.06 | 1.04 | 1.31 | 1.08 | 0.84 | 0.55 | 0.74 | 0.85 | 0.80 | 0.73 | -0.13 | 0.42 |
Upper | 1.84 | 1.47 | 0.91 | 1.00 | 1.06 | 1.05 | 1.31 | 1.09 | 0.86 | 0.61 | 0.77 | 0.88 | 0.81 | 0.75 | -0.09 | 0.43 |
Mean | 1.88 | 1.58 | 0.94 | 1.00 | 1.12 | 1.07 | 1.32 | 1.12 | 0.95 | 0.69 | 0.81 | 0.93 | 0.80 | 0.77 | 0.08 | 0.41 |
SD | 0.039 | 0.001 | 0.002 | 0.02 | 0.001 | 0.017 | 0.007 | 0.002 | 0.014 | 0.003 | 0.004 | 0.007 | 0.006 | 0.022 | 0.016 | 0.01 |
Lower | 1.85 | 1.58 | 0.93 | 0.98 | 1.12 | 1.05 | 1.31 | 1.12 | 0.94 | 0.69 | 0.80 | 0.92 | 0.79 | 0.74 | 0.07 | 0.40 |
Upper | 1.92 | 1.58 | 0.94 | 1.02 | 1.12 | 1.09 | 1.33 | 1.12 | 0.96 | 0.69 | 0.81 | 0.94 | 0.80 | 0.78 | 0.10 | 0.42 |
Value | 1.87 | 1.50 | 0.87 | 1.02 | 1.13 | 1.07 | 1.33 | 1.11 | 0.91 | 0.72 | 0.79 | 0.94 | 0.83 | 0.75 | 0.51 | 0.43 |
Value | 1.89 | 1.59 | 0.96 | 1.03 | 1.13 | 1.12 | 1.34 | 1.12 | 0.95 | 0.85 | 0.76 | 0.96 | 0.82 | 0.73 | 0.28 | 0.49 |
Mean | 1.92 | 1.59 | 1.01 | 1.02 | 1.17 | 1.1 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
SD | 0.035 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 |
Lower | 1.89 | 1.59 | 1.01 | 1.02 | 1.17 | 1.10 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
Upper | 1.94 | 1.59 | 1.01 | 1.02 | 1.17 | 1.10 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
Mean | 1.85 | 1.53 | 0.89 | 0.98 | 1.04 | 1.03 | 1.30 | 1.16 | 0.91 | 0.76 | 0.80 | 0.91 | 0.77 | 0.75 | 0.05 | 0.34 |
SD | 0.025 | 0.01 | 0.002 | 0.007 | 0.007 | 0.005 | 0.005 | 0.001 | 0.013 | 0.007 | 0.024 | 0.005 | 0.004 | 0.025 | 0.019 | 0.016 |
Lower | 1.83 | 1.53 | 0.89 | 0.97 | 1.03 | 1.03 | 1.29 | 1.16 | 0.91 | 0.75 | 0.79 | 0.91 | 0.77 | 0.74 | 0.04 | 0.33 |
Upper | 1.87 | 1.54 | 0.90 | 0.98 | 1.04 | 1.04 | 1.30 | 1.16 | 0.92 | 0.76 | 0.82 | 0.91 | 0.78 | 0.77 | 0.06 | 0.35 |
Mean | 1.85 | 1.49 | 1.01 | 0.95 | 1.05 | 1.02 | 1.30 | 1.14 | 0.92 | 0.71 | 0.75 | 0.91 | 0.77 | 0.67 | 0.09 | 0.35 |
SD | 0.026 | 0.028 | 0.055 | 0.029 | 0.027 | 0.023 | 0.009 | 0.018 | 0.025 | 0.036 | 0.034 | 0.014 | 0.015 | 0.087 | 0.069 | 0.022 |
Lower | 1.79 | 1.45 | 0.91 | 0.90 | 0.99 | 0.97 | 1.29 | 1.10 | 0.87 | 0.62 | 0.70 | 0.89 | 0.74 | 0.46 | 0.00 | 0.32 |
Upper | 1.88 | 1.54 | 1.08 | 1.00 | 1.09 | 1.05 | 1.32 | 1.16 | 0.95 | 0.75 | 0.80 | 0.92 | 0.79 | 0.73 | 0.23 | 0.38 |
Value | 1.76 | 1.80 | 0.95 | 0.99 | 1.09 | 1.02 | 1.28 | 1.10 | 0.76 | 0.72 | 0.74 | 0.88 | 0.75 | 0.68 | 0.19 | 0.32 |
Mean | 1.86 | 1.53 | 0.91 | 0.97 | 1.08 | 1.02 | 1.30 | 1.16 | 0.89 | 0.71 | 0.77 | 0.90 | 0.77 | 0.74 | 0.09 | 0.34 |
SD | 0.024 | 0.016 | 0.034 | 0.021 | 0.032 | 0.019 | 0.014 | 0.017 | 0.008 | 0.015 | 0.004 | 0.012 | 0.007 | 0.012 | 0.061 | 0.023 |
Lower | 1.81 | 1.50 | 0.86 | 0.93 | 1.02 | 0.99 | 1.27 | 1.14 | 0.88 | 0.69 | 0.76 | 0.89 | 0.76 | 0.72 | 0.00 | 0.30 |
Upper | 1.88 | 1.55 | 0.95 | 1.00 | 1.12 | 1.05 | 1.32 | 1.19 | 0.90 | 0.74 | 0.78 | 0.93 | 0.78 | 0.76 | 0.18 | 0.38 |
Value | 1.83 | 1.53 | 1.01 | 0.99 | 1.13 | 1.09 | 1.30 | 1.14 | 0.75 | 0.70 | 0.78 | 0.92 | 0.78 | 0.61 | 0.08 | 0.35 |
Mean | 1.83 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
SD | 0.040 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 |
Lower | 1.80 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
Upper | 1.86 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
Adult female ZMKU R 00947 from Pa La-U Waterfall, Kaeng Krachan National Park, Huai Sat Yai Subdistrict, Hua Hin District, Prachuap Khiri Khan Province, Thailand (
Adult female ZMKU R 00946 bearing the same data as the holotype.
Holotype of
Body relatively short (
Forelimbs moderate in stature, relatively short (
Posterior one-half of tail regenerated, tail long 91.5 mm (
Type series of
The specific epithet
The holotype and paratype were collected at night (1900–2055 h) on granite boulders by a rocky stream dry evergreen forest at 368 m in elevation (Fig.
Habitat of
Sex and raw meristic, categorical, and morphometric data used in the analyses from specimens in the
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Institutional catalog number | LSUHC 1899 | LSUHC 15076 | LSUHC 11788 | THNHM 10670 | THNHM 14112 | LSUHC 8238 | LSUHC 12180 | LSUHC 12181 | ZMMU R-16144 | THNHM 20226 paratype | THNHM 20228 paratype | THNHM 20229 paratype | THNHM 20227 paratype | ZMMU R-16492 | HNUE-R00111 holotype | IEBR 4829 paratype | VNUF R.2020.12 paratype | HNUE-R00112 paratype | FMNH 255454 | FMNH 270493 | FMNH 270492 | FMNH 265806 | NCSM 79472 | ZMMU R-14917 | NCSM 80100 | ZMMU R-16851 holotype | ZMMU R-16852 paratype | ZMKU R 00948 topotype | THNHM 24622 | THNHM 24838 | THNHM 03251 | THNHM 03252 | THNHM 03253 | THNHM 03254 | THNHM 01807 | ZMKU R 00959 holotype | ZMKU R 00958 paratype | ZMKU R 00960 paratype | ZMKU R 00961 paratype | ZMKU R 00962 paratype | ZMKU R 00963 paratype | ZMKU R 00964 paratype | ZMKU R 00945 holotype | THNHM 01667 | ZMKU R 00949 holotype | ZMKU R 00947 holotype | ZMKU R 00946 paratype | ZMKU R 00950 | ZMKU R 00951 | ZMKU R 00952 | ZMKU R 00953 | ZMKU R 00954 | ZMKU R 00955 | ZMKU R 00956 | ZMKU R 00957 | THNHM 00104 | THNHM 27821 |
Sex | m | f | f | f | f | f | m | m | f | f | f | f | f | m | m | f | f | f | f | m | m | m | f | f | f | m | f | f | m | f | m | m | f | m | m | m | m | m | f | f | f | f | f | m | m | f | f | f | m | f | m | m | f | m | f | f | f |
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Supralabials ( |
11 | 12 | 10 | 14 | 12 | 11 | 8 | 13 | 9 | 14 | 12 | 11 | 12 | 11 | 10 | 10 | 10 | 10 | 13 | 13 | 13 | 10 | 14 | 9 | 12 | 11 | 9 | 14 | 11 | 13 | 13 | 11 | 12 | 13 | 12 | 12R/12L | 13R/12L | 13R/12L | 11R/12L | 12R/12L | 13R/12L | 12R/11L | 12R/13L | 12 | 13R/15L | 13R/12L | 13R/12L | 12 | 13 | 13 | 14 | 13 | 13 | 13 | 13 | 12 | 15 |
Infralabials ( |
8 | 10 | 9 | 11 | 11 | 11 | 8 | 11 | 9 | 9 | 8 | 8 | 7 | 9 | 9 | 9 | 9 | 9 | 10 | 9 | 11 | 8 | 11 | 10 | 12 | 10 | 11 | 9 | 10 | 10 | 10 | 10 | 10 | 11 | 10 | 10R/10L | 10R/10L | 9R/10L | 10R/10L | 10R/10L | 10R/10L | 10R/10L | 9R/9L | 10 | 10R/11L | 11R/10L | 10R/9L | 8 | 8 | 10 | 10 | 9 | 10 | 10 | 9 | 10 | 11 |
Paravertebral tubercles ( |
39 | 37 | 38 | 37 | 37 | 0 | 0 | 0 | 0 | 32 | 33 | 33 | 33 | 30 | 39 | 40 | 38 | 40 | 28 | 27 | 26 | 27 | 28 | 32 | 29 | 27 | 30 | 30 | 26 | 28 | 27 | 27 | 30 | 30 | 26 | 30 | 28 | 27 | 27 | 28 | 26 | 28 | 34 | 29 | 33 | 34 | 33 | 32 | 33 | 34 | 34 | 36 | 36 | 30 | 30 | 33 | 29 |
Longitudinal rows of tubercles ( |
15 | 16 | 17 | 16 | 14 | 6 | 7 | 4 | 4 | 19 | 20 | 19 | 19 | 18 | 18 | 18 | 17 | 22 | 19 | 18 | 17 | 19 | 18 | 24 | 19 | 19 | 20 | 19 | 18 | 19 | 18 | 18 | 19 | 19 | 19 | 17 | 17 | 14 | 16 | 17 | 18 | 16 | 14 | 19 | 17 | 20 | 18 | 21 | 19 | 20 | 20 | 21 | 21 | 19 | 19 | 18 | 20 |
Ventral scales ( |
38 | 38 | 38 | 36 | 39 | 45 | 45 | 47 | 36 | 42 | 40 | 42 | 43 | 34 | 38 | 36 | 35 | 32 | 37 | 36 | 36 | 33 | 33 | 36 | 35 | 34 | 43 | 38 | 38 | 36 | 37 | 37 | 39 | 34 | 35 | 34 | 37 | 33 | 30 | 36 | 37 | 39 | 35 | 34 | 36 | 34 | 37 | 34 | 33 | 33 | 34 | 30 | 33 | 32 | 33 | 37 | 36 |
Ventral scales along middle of the body ( |
176 | 170 | 182 | 154 | 160 | 190 | 225 | 234 | 192 | 187 | 170 | 187 | 178 | 160 | 168 | 164 | 178 | 158 | 159 | 166 | 156 | 158 | 164 | 166 | 165 | 154 | 152 | 165 | 162 | 158 | 157 | 159 | 168 | 160 | 161 | 155 | 154 | 155 | 172 | 164 | 175 | 170 | 172 | 159 | 159 | 160 | 166 | 173 | 158 | 156 | 166 | 159 | 159 | 150 | 169 | 159 | 165 |
Expanded subdigital lamellae on 4th toe ( |
7 | 8 | 9 | 8 | 8 | 10 | 9 | 9 | 9 | 12 | 10 | 10 | 11 | 9 | 8 | 10 | 9 | 9 | 10 | 10 | 8 | 10 | 9 | 8 | 10 | 9 | 9 | 9 | 8 | 9 | 9 | 10 | 9 | 10 | 10 | 9R/9L | 10R/10L | 9R/9L | 9R/9L | 10R/11L | 9R/10L | 9R/9L | 9R/8L | 8 | 8R/(broken)L | 9R/9L | 9R/9L | 9 | 10 | 9 | 8 | 10 | 8 | 9 | 9 | 9 | 7 |
Unmodified subdigital lamellae on 4th toe ( |
13 | 11 | 11 | 11 | 12 | 11 | 10 | 11 | 9 | 14 | 13 | 12 | 14 | 10 | 11 | 10 | 11 | 10 | 11 | 11 | 11 | 11 | 12 | 10 | 10 | 11 | 11 | 12 | 11 | 13 | 12 | 12 | 15 | 13 | 13 | 11R/11L | 12R/11L | 10R/10L | 12R/12L | 11R/11L | 10R/10L | 12R/13L | 12R/11L | 13 | 12R/(broken)L | 13R/13L | 12R/13L | 12 | 14 | 13 | 12 | 13 | 12 | 11 | 13 | 12 | 12 |
Total subdigital lamellae 4th toe ( |
20 | 19 | 20 | 19 | 20 | 21 | 19 | 20 | 18 | 26 | 23 | 22 | 23 | 19 | 13 | 16 | 17 | 16 | 21 | 21 | 19 | 21 | 21 | 18 | 20 | 20 | 18 | 21 | 19 | 22 | 21 | 22 | 14 | 23 | 23 | 20R/20L | 22R/21L | 19R/19L | 21R/21L | 21R/22L | 19R/20L | 22R/22L | 21R/19L | 21 | 20 | 22R/22L | 21R/22L | 21 | 24 | 22 | 20 | 23 | 20 | 20 | 22 | 21 | 19 |
Expanded subdigital lamellae on 4th finger ( |
8 | 8 | 8 | 7 | 8 | 8 | 9 | 9 | 9 | 9 | 8 | 9 | 9 | 10 | 6 | 6 | 7 | 6 | 8 | 8 | 8 | 8 | 9 | 7 | 9 | 9 | 8 | 8 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8R/8L | 8R/8L | 8R/8L | 8R/8L | 7R/7L | 8R/9L | 7R/7L | 8R/8L | 8 | 7R/7L | 8R/8L | 8R/8L | 8 | 7 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
Unmodified subdigital lamellae on 4th finger ( |
9 | 11 | 10 | 10 | 10 | 12 | 13 | 9 | 8 | 12 | 11 | 12 | 13 | 9 | 9 | 9 | 9 | 9 | 10 | 10 | 10 | 10 | 8 | 9 | 10 | 10 | 9 | 11 | 10 | 11 | 10 | 10 | 12 | 12 | 12 | 10R/10L | 10R/10L | 10R/9L | 11R/11L | 10R/10L | 9R/9L | 10R/10L | 10R/10L | 12 | 11R/11L | 11R/10L | 12R/12L | 10 | 12 | 12 | 11 | 12 | 12 | 11 | 12 | 11 | 10 |
Total subdigital lamellae 4th finger ( |
17 | 19 | 18 | 17 | 18 | 20 | 22 | 18 | 17 | 21 | 21 | 21 | 22 | 19 | 15 | 15 | 18 | 15 | 18 | 18 | 18 | 18 | 17 | 16 | 19 | 19 | 17 | 19 | 17 | 17 | 18 | 18 | 20 | 20 | 20 | 18R/18L | 18R/18L | 18R/17L | 19R/19L | 17R/17L | 17R/18L | 17R/17L | 18R/18L | 20 | 18R/18L | 19R/18L | 20R/20L | 18 | 19 | 19 | 19 | 20 | 20 | 19 | 20 | 19 | 18 |
Enlarged femoral scales (R/L) | 0 | 0 | 0 | 8R/8L | 7R/7L | 0 | 0 | 0 | 0 | 11R/8L | 10R/9L | 8R/8L | 9R/10L | 9R/8L | 10R/10L | 9R/8L | 10R/9L | 8R/9L | 9R/7L | 8R/9L | 9R/9L | 8R/8L | 9R/8L | 7R/8L | 7R/8L | 9R/8L | 8R/8L | 9R/8L | 9R/L | 9R/9L | 9R/7L | 7R/7L | 6R/7L | 5R/8L | 7R/7L | 5R/6L | 4R/5L | 5R/6L | 6R/6L | 5R/6L | 5R/6L | 6R/6L | 6R/6L | 7R/7L | 8R/8L | 8R/8L | 6R/8L | 5R/7L | 8R/8L | 8R/8L | 7R/8L | 8R/8L | 7R/8L | 7R/6L | 8R/8L | 9R/9L | 7R/10L |
Total enlarged femoral scales ( |
16 | 10 | 11 | 16 | 14 | 0 | 0 | 0 | 0 | 14 | 19 | 19 | 19 | 17 | 20 | 17 | 19 | 17 | 16 | 17 | 18 | 16 | 17 | 15 | 15 | 17 | 16 | 17 | 18 | 18 | 16 | 14 | 13 | 13 | 14 | 11 | 9 | 11 | 12 | 11 | 11 | 12 | 12 | 14 | 16 | 16 | 14 | 12 | 16 | 16 | 15 | 16 | 15 | 13 | 16 | 18 | 17 |
Total femoral pores in males ( |
7 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 14 | 0 | 0 | 0 | 0 | 14 | 15 | 13 | 0 | 0 | 0 | 17 | 0 | 0 | 14 | 0 | 12 | 13 | 0 | 11 | 13 | 11 | 8 | 10 | 0 | 0 | 0 | 0 | 0 | 14 | 12 | 0 | 0 | 0 | 16 | 0 | 14 | 15 | 14 | 12 | 0 | 0 | 0 |
Enlarged precloacal scales ( |
7 | 7 | 7 | 8 | 7 | 8 | 8 | 8 | 7 | 14 | 15 | 13 | 19 | 13 | 13 | 13 | 13 | 13 | 15 | 13 | 13 | 13 | 12 | 13 | 13 | 17 | 13 | 15 | 15 | 15 | 14 | 13 | 15 | 15 | 14 | 15 | 14 | 14 | 15 | 14 | 15 | 15 | 12 | 16 | 14 | 15 | 15 | 17 | 15 | 15 | 15 | 15 | 15 | 15 | 15 | 14 | 16 |
Precloacal pores in males ( |
7 | 0 | 0 | 0 | 0 | 0 | 8 | 8 | 0 | 0 | 0 | 0 | 0 | 13 | 0 | 0 | 0 | 0 | 13 | 13 | 13 | 13 | 0 | 0 | 0 | 17 | 13 | 0 | 15 | 0 | 14 | 13 | 0 | 15 | 14 | 15 | 14 | 14 | 0 | 0 | 0 | 0 | 0 | 16 | 14 | 0 | 0 | 0 | 15 | 0 | 15 | 15 | 15 | 15 | 0 | 0 | 0 |
Postcloacal tubercles (PCT) | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 3 | 2 | 1 | 2 | 0 | 0 | 0 | 0 | 2 | 3 | 4 | 3 | 2 | 2R/3L | 3 | 2 | 3 | 2 | 2 | 3 | 2 | 3R/2L | 3R/2L | 3R/3L | 1R/1L | 3R/2L | 3R/3L | 2R/2L | 1R/1L | 3 | 3R/3L | 2R/2L | 3R/3L | 2 | 2R/3L | 3 | 3 | 2R/3L | 2R/3L | 3 | 2 | 3 | 3 |
Body bands ( |
4 | 6 | 3 | 5 | 5 | 5 | 5 | 3 | 3 | 5 | 5 | 5 | 5 | 3 | 6 | 6 | 6 | 6 | 3 | 4 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 4 | 4 | / | / | 5 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 5 | 5 | 6 | 3 | 4 | 3 | 4 | 3 | 4 | 3 | 5 | 4 | 4 | 3 | / |
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Small tubercles on flank ( |
present | present | present | present | present | absent | absent | absent | absent | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present |
Dorsolateral caudal tubercles ( |
small | small | small | / | small | large | large | large | large | small | / | small | small | large | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | / | small | small | small | small | small | small | small | large | large | large | large | large | large | large | large | large | large | large | / | large | small | small |
Ventrolateral caudal fringe narrow or wide ( |
narrow | narrow | narrow | / | narrow | wide | wide | wide | wide | narrow | / | narrow | narrow | wide | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | / | narrow | narrow | narrow | narrow | narrow | narrow | narrow | wide | wide | wide | wide | wide | wide | wide | wide | wide | wide | wide | / | wide | narrow | narrow |
Ventrolateral caudal fringe scales generally homogenous ( |
no | no | no | / | no | no | no | no | no | yes | yes | yes | yes | yes | no | no | no | no | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | / | no | no | no | no | no | no | no | no | no | no | yes | yes | no | no | no | no | no | no | / | no | yes | yes |
Tail cross-section ( |
circular | circular | circular | / | circular | square | square | square | square | circular | / | circular | circular | square | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | square | square | square | square | square | square | square | square | square | / | circular | circular | circular | circular | circular | circular | circular | square | / | circular | square | square | square | square | square | square | square | square | / | square | circular | circular |
Slightly enlarged medial subcaudals ( |
present | present | present | / | absent | absent | absent | absent | absent | absent | / | absent | absent | present | present | present | present | present | / | present | present | present | present | present | present | absent | absent | absent | absent | absent | absent | absent | absent | absent | / | present | present | present | present | present | present | present | present | present | present | absent | absent | present | present | present | present | present | present | / | present | present | present |
Single enlarged medial subcaudal ( |
absent | absent | absent | / | absent | absent | absent | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | present | present | present | present | present | present | present | present | present | / | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | present | present | absent | absent | absent | absent | absent | absent | / | absent | absent | absent |
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate ( |
no | no | no | / | no | no | no | no | no | yes | / | yes | yes | no | no | no | no | no | / | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | yes | no | no | no | no | no | no | yes | no | / | no | no | no |
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68.8 | 70.8 | 64.1 | 65.95 | 63.79 | 80.2 | 78.2 | 84.8 | 78.6 | 81.19 | 74.80 | 78.56 | 59.70 | 68.1 | 66.5 | 68.1 | 69.3 | 46.6 | 83.6 | 70.2 | 74.1 | 73.8 | 78.0 | 87.1 | 77.7 | 74.9 | 71.7 | 71.6 | 68.3 | 71.8 | 73.6 | 75.3 | 74.7 | 73.2 | 61.5 | 72.5 | 72.0 | 69.6 | 68.4 | 76.8 | 65.7 | 78.2 | 60.1 | 70.2 | 58.1 | 73.9 | 68.1 | 73.1 | 73.5 | 73.7 | 73.2 | 64.4 | 76.6 | 76.6 | 74.2 | 63.7 | 72.9 |
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35.7 | 33.4 | 30.1 | 30.0 | 26.5 | 39.7 | 37.8 | 41.5 | 36.2 | 34.5 | 33.7 | 32.7 | 24.6 | 34.6 | 28.8 | 29.8 | 30.2 | 19.7 | 41.3 | 35.4 | 37.0 | 31.3 | 38.2 | 41.9 | 36.8 | 34.6 | 32.6 | 33.9 | 27.3 | 29.9 | 30.9 | 31.3 | 32.2 | 30.3 | 26.2 | 33.4 | 33.6 | 32.0 | 30.4 | 35.6 | 30.6 | 38.1 | 29.0 | 31.5 | 26.6 | 34.8 | 33.2 | 34.8 | 33.9 | 35.4 | 33.6 | 28.5 | 37.1 | 33.2 | 35.1 | 25.8 | 30.6 |
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9.7 | 9.3 | 8.0 | 9.6 | 9.5 | 10.2 | 9.1 | 10.1 | 1.7 | 9.8 | 10.2 | 11.2 | 7.4 | 10.3 | 7.9 | 8.1 | 8.5 | 5.6 | 8.6 | 8.7 | 8.6 | 6.9 | 8.7 | 11.5 | 9.2 | 10.7 | 10.4 | 7.9 | 9.8 | 8.3 | 12.2 | 11.3 | 11.8 | 11.0 | 10.1 | 9.1 | 8.8 | 9.0 | 8.0 | 10.0 | 7.5 | 10.1 | 6.5 | 10.2 | 7.0 | 8.1 | 7.6 | 8.4 | 7.2 | 9.0 | 9.0 | 7.2 | 8.0 | 8.1 | 8.6 | 7.6 | 10.1 |
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9.9 | 9.8 | 8.9 | 8.2 | 8.7 | 11.5 | 11.7 | 11.8 | 10.2 | 10.6 | 10.5 | 11.1 | 8.4 | 8.5 | 9.2 | 10.0 | 10.1 | 6.5 | 10.2 | 9.3 | 10.4 | 10.0 | 10.3 | 10.4 | 10.7 | 8.6 | 7.9 | 9.6 | 8.7 | 8.5 | 9.0 | 10.6 | 9.6 | 9.2 | 7.9 | 10.5 | 10.3 | 10.5 | 10.1 | 11.1 | 8.8 | 10.8 | 7.6 | 8.6 | 8.3 | 9.7 | 9.1 | 9.5 | 9.1 | 9.2 | 9.8 | 9.2 | 10.0 | 8.6 | 9.8 | 8.1 | 9.6 |
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12.0 | 12.6 | 11.5 | 11.7 | 9.8 | 12.9 | 14.2 | 14.6 | 13.1 | 14.7 | 13.2 | 12.7 | 10.2 | 12.6 | 11.5 | 11.5 | 11.5 | 7.6 | 13.7 | 12.7 | 13.0 | 13.1 | 13.1 | 15.2 | 14.2 | 12.6 | 11.8 | 10.5 | 10.8 | 10.9 | 11.5 | 10.2 | 11.9 | 12.1 | 9.5 | 13.1 | 12.5 | 12.5 | 13.5 | 14.1 | 11.5 | 13.9 | 10.4 | 12.1 | 10.0 | 11.4 | 10.4 | 12.8 | 11.6 | 12.3 | 12.5 | 10.9 | 13.7 | 10.8 | 12.5 | 10.7 | 12.8 |
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11.6 | 12.2 | 10.5 | 9.7 | 8.2 | 13.5 | 14.0 | 13.8 | 12.3 | 13.1 | 11.9 | 12.9 | 10.2 | 11.4 | 10.8 | 11.1 | 11.8 | 7.8 | 12.5 | 11.8 | 11.2 | 11.1 | 12.8 | 12.6 | 12.7 | 10.1 | 9.3 | 11.2 | 9.7 | 10.7 | 10.9 | 11.7 | 11.3 | 11.1 | 9.1 | 11.3 | 10.6 | 10.2 | 9.9 | 11.2 | 9.4 | 12.3 | 8.4 | 11.8 | 8.4 | 11.2 | 10.3 | 10.5 | 10.1 | 10.6 | 10.6 | 9.9 | 11.1 | 10.0 | 11.4 | 10.1 | 10.2 |
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19.3 | 19.3 | 19.0 | 17.9 | 18.2 | 21.8 | 21.6 | 21.9 | 21.7 | 20.8 | 19.9 | 21.7 | 16.7 | 18.4 | 20.1 | 20.4 | 20.7 | 16.1 | 21.7 | 20.6 | 20.3 | 20.7 | 21.2 | 22.1 | 21.4 | 20.2 | 19.2 | 19.7 | 19.7 | 19.9 | 20.8 | 21.3 | 20.8 | 21.5 | 17.9 | 20.1 | 20.5 | 19.7 | 20.1 | 21.2 | 18.6 | 21.3 | 17.3 | 18.3 | 16.1 | 20.3 | 19.3 | 19.9 | 20.9 | 20.1 | 20.0 | 17.6 | 20.4 | 19.3 | 20.0 | 17.6 | 19.9 |
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13.2 | 13.8 | 12.3 | 12.3 | 12.0 | 15.6 | 16.1 | 15.9 | 15.1 | 14.0 | 13.4 | 14.2 | 11.4 | 13.1 | 12.6 | 12.0 | 11.8 | 8.8 | 13.8 | 12.5 | 13.0 | 12.3 | 12.7 | 14.8 | 13.5 | 14.6 | 13.4 | 14.0 | 13.1 | 13.9 | 14.9 | 15.0 | 13.1 | 14.1 | 11.8 | 14.0 | 13.4 | 12.9 | 13.0 | 14.9 | 13.0 | 15.4 | 11.6 | 12.1 | 10.9 | 14.9 | 13.7 | 14.5 | 14.3 | 15.7 | 13.9 | 12.8 | 14.7 | 14.4 | 14.1 | 11.9 | 13.8 |
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8.0 | 7.6 | 7.6 | 7.3 | 7.0 | 9.6 | 9.8 | 10.4 | 9.8 | 3.4 | 8.6 | 8.7 | 6.6 | 8.3 | 7.4 | 7.2 | 6.6 | 5.1 | 9.2 | 8.4 | 9.1 | 7.6 | 8.3 | 8.7 | 9.2 | 9.2 | 8.5 | 8.3 | 7.3 | 8.9 | 8.2 | 8.2 | 8.1 | 8.9 | 7.5 | 8.5 | 8.1 | 8.3 | 7.9 | 8.1 | 7.8 | 8.3 | 6.5 | 7.8 | 6.3 | 8.2 | 8.2 | 7.8 | 7.7 | 7.9 | 7.7 | 7.0 | 8.2 | 7.8 | 7.6 | 7.7 | 8.4 |
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5.2 | 4.5 | 4.3 | 5.3 | 4.4 | 4.8 | 5.0 | 5.7 | 5.0 | 5.3 | 5.5 | 5.9 | 4.4 | 4.4 | 3.8 | 4.1 | 3.4 | 2.6 | 4.9 | 4.9 | 4.9 | 4.8 | 6.5 | 4.6 | 6.0 | 4.6 | 4.3 | 5.5 | 4.9 | 5.1 | 5.8 | 5.4 | 5.0 | 5.5 | 4.7 | 5.0 | 5.0 | 4.9 | 4.7 | 5.1 | 4.5 | 5.3 | 4.2 | 5.2 | 4.6 | 5.8 | 5.6 | 5.0 | 5.1 | 5.0 | 5.0 | 4.8 | 5.6 | 5.3 | 4.9 | 4.1 | 5.3 |
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5.7 | 5.9 | 4.9 | 5.7 | 5.7 | 6.4 | 7.1 | 7.0 | 6.8 | 5.8 | 6.2 | 6.4 | 4.8 | 6.2 | 5.8 | 5.5 | 5.9 | 4.4 | 6.9 | 6.1 | 6.2 | 5.7 | 5.3 | 6.5 | 6.2 | 6.2 | 6.2 | 5.8 | 5.1 | 6.2 | 5.6 | 5.7 | 5.4 | 6.2 | 4.3 | 6.5 | 5.9 | 5.7 | 5.8 | 6.1 | 5.4 | 6.5 | 5.0 | 4.9 | 4.7 | 6.5 | 6.2 | 5.9 | 5.9 | 6.0 | 5.9 | 5.3 | 6.1 | 6.0 | 6.0 | 4.9 | 6.3 |
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7.4 | 7.6 | 7.0 | 7.0 | 7.2 | 8.6 | 8.7 | 9.5 | 8.6 | 8.3 | 7.8 | 9.1 | 6.8 | 7.7 | 7.5 | 7.6 | 6.9 | 5.0 | 9.0 | 8.3 | 8.3 | 8.2 | 8.7 | 8.8 | 8.4 | 8.3 | 7.7 | 7.9 | 7.4 | 8.1 | 8.4 | 8.8 | 8.1 | 8.6 | 7.3 | 8.5 | 8.3 | 8.2 | 8.1 | 9.2 | 7.3 | 9.3 | 6.9 | 7.5 | 6.4 | 8.3 | 7.9 | 7.9 | 8.5 | 7.9 | 7.9 | 7.3 | 8.2 | 7.9 | 7.9 | 7.2 | 8.0 |
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5.7 | 5.4 | 4.9 | 5.3 | 5.4 | 6.0 | 6.2 | 6.5 | 6.2 | 6.0 | 5.5 | 6.8 | 5.1 | 5.5 | 6.7 | 6.3 | 6.2 | 4.5 | 6.5 | 6.2 | 6.1 | 6.2 | 6.2 | 6.6 | 6.0 | 6.3 | 5.7 | 5.8 | 5.4 | 6.0 | 6.2 | 6.4 | 5.8 | 6.2 | 5.3 | 6.5 | 6.2 | 5.9 | 6.1 | 6.6 | 5.6 | 6.5 | 5.2 | 5.5 | 4.9 | 6.1 | 5.8 | 6.0 | 6.1 | 6.0 | 5.8 | 5.4 | 6.1 | 6.0 | 5.9 | 5.6 | 5.9 |
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5.4 | 4.7 | 4.7 | 4.2 | 5.2 | 5.7 | 5.4 | 5.4 | 3.9 | 4.8 | 4.7 | 5.5 | 4.3 | 2.9 | 5.6 | 5.4 | 5.6 | 4.2 | 6.6 | 5.6 | 5.4 | 5.1 | 4.9 | 3.5 | 5.7 | 3.3 | 3.1 | 5.6 | 4.5 | 4.7 | 5.6 | 5.7 | 5.7 | 5.6 | 4.2 | 5.5 | 5.4 | 5.3 | 5.1 | 5.6 | 5.0 | 5.6 | 4.2 | 4.0 | 4.3 | 5.8 | 5.5 | 5.4 | 5.5 | 5.8 | 5.5 | 4.9 | 5.7 | 5.6 | 5.3 | 4.8 | 6.1 |
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1.0 | 1.4 | 1.1 | 1.3 | 1.0 | 1.9 | 1.4 | 1.5 | 1.4 | 1.3 | 1.3 | 1.6 | 1.2 | 0.9 | 0.8 | 0.8 | 0.7 | 0.3 | 1.3 | 1.1 | 1.2 | 1.0 | 1.5 | 1.2 | 0.9 | 1.2 | 1.0 | 1.4 | 1.6 | 1.5 | 1.2 | 1.3 | 1.2 | 1.2 | 0.9 | 1.4 | 1.5 | 1.7 | 1.4 | 1.8 | 1.6 | 1.8 | 1.0 | 1.3 | 1.5 | 1.1 | 1.1 | 1.1 | 1.5 | 1.5 | 1.2 | 1.2 | 1.0 | 1.2 | 1.3 | 1.4 | 1.4 |
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1.7 | 2.1 | 2.3 | 2.1 | 2.2 | 2.7 | 2.6 | 2.5 | 3.1 | 2.1 | 2.2 | 2.5 | 1.8 | 2.3 | 2.8 | 2.6 | 2.6 | 2.0 | 2.8 | 2.5 | 2.5 | 2.3 | 2.7 | 2.2 | 2.5 | 2.2 | 2.1 | 2.1 | 2.0 | 2.2 | 2.4 | 2.5 | 2.4 | 2.3 | 2.0 | 2.3 | 2.4 | 2.5 | 2.3 | 2.3 | 2.3 | 2.6 | 1.9 | 2.2 | 1.8 | 2.3 | 2.0 | 2.3 | 2.4 | 2.2 | 2.0 | 2.0 | 2.3 | 2.2 | 2.2 | 2.1 | 2.3 |
Adult female ZMKU R 00945 from Ko Chang Island, Ko Phayam Subdistrict, Mueang Ranong District, Ranong Province, Thailand (
Holotype of
Body relatively short (
Forelimbs moderate in stature, relatively short (
Tail original (but in two pieces), long 80.3 mm (
The additional specimen (THNHM 01667) closely approximates the holotype in overall coloration and pattern except that it is more boldly marked. It has four dark-colored body bands as opposed to three and a complete original tail bearing eight dark-colored and seven pale-colored bands. The pale-colored postorbital stripe is slightly thinner and less distinct. Meristic and morphometric differences are listed in Table
The specific epithet
The holotype (ZMKU R 00945) was collected at night (2107 h) among branches of a small tree approximately 100 cm above the ground at 36 m elevation with a temperature of 28.6 °C and relative humidity of 83.9%. The surrounding habitat was dry evergreen forest with a rocky stream nearby (Fig.
Habitat of
Adult male ZMKU R 00949 from Thung Na Ngam Subdistrict, Lan Sak District, Uthai Thani Province, Thailand (
Significant
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0.00 | 0.01 | < 0.001 | 0.001 | 0.048 | < 0.001 | 0.004 | 0.002 | 0.004 | |||||||
0.00 | 0.00 | 0.00 | < 0.001 | ||||||||||||
0.00 | 0.003 | 0.001 | < 0.001 | ||||||||||||
0.014 | 0.03 | 0.01 | < 0.001 | 0.008 | 0.038 | < 0.001 | 0.004 | 0.000 | |||||||
< 0.001 | 0.001 | 0.022 | < 0.001 | 0.021 | |||||||||||
0.01 | < 0.001 | < 0.001 | 0.03 | ||||||||||||
0.01 | |||||||||||||||
< 0.001 | 0.002 | 0.04 | < 0.001 | 0.005 | < 0.001 | < 0.001 | |||||||||
< 0.001 | 0.025 | 0.038 | |||||||||||||
0.049 | < 0.001 | < 0.001 | 0.004 | ||||||||||||
0.0 | 0.007 | 0.013 | 0.023 | 0.007 | |||||||||||
0.007 | 0.031 | 0.010 | < 0.001 | 0.000 | |||||||||||
0.011 | 0.003 | ||||||||||||||
0.0 | 0.01 | 0.00 | 0.044 | ||||||||||||
0.00 | 0.007 | ||||||||||||||
< 0.001 | 0.019 | 0.006 | < 0.001 | 0.001 | |||||||||||
0.003 | < 0.001 | < 0.001 | 0.046 | < 0.001 | 0.001 | < 0.001 | < 0.001 | 0.001 | |||||||
< 0.001 | < 0.001 | 0.042 | 0.007 | < 0.001 | < 0.001 | 0.000 | |||||||||
< 0.001 | 0.021 | 0.0 | 0.00 | 0.007 | 0.02 | 0.006 | |||||||||
0.001 | 0.01 | 0.03 | 0.003 | 0.043 | 0.001 | 0.019 | 0.019 | 0.003 | |||||||
0.02 | 0.004 | 0.02 | 0.033 | ||||||||||||
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0.037 | < 0.001 | 0.05 | |||||||||||||
< 0.001 | 0.000 | 0.002 | 0.044 | ||||||||||||
< 0.001 | 0.028 | < 0.001 | |||||||||||||
0.017 | 0.029 | ||||||||||||||
< 0.001 | < 0.001 | ||||||||||||||
< 0.001 | 0.003 | 0.022 | < 0.001 | 0.05 | |||||||||||
0.000 | 0.000 | 0.021 | < 0.001 | 0.000 | < 0.001 | ||||||||||
0.037 | 0.002 | < 0.001 | < 0.001 | 0.033 | 0.000 | 0.002 | < 0.001 | ||||||||
0.005 | |||||||||||||||
0.000 | 0.001 | 0.001 | |||||||||||||
< 0.001 | < 0.001 | 0.001 | 0.004 | 0.020 | |||||||||||
0.001 | 0.029 | < 0.001 | < 0.001 | 0.029 | |||||||||||
0.029 | 0.025 | 0.005 | |||||||||||||
0.004 | 0.011 | 0.001 | 0.044 | 0.001 | 0.002 | ||||||||||
0.003 | < 0.001 | 0.005 | 0.043 | 0.01 | < 0.001 | ||||||||||
0.0267 | 0.001 | < 0.001 | |||||||||||||
< 0.001 | < 0.001 | 0.008 | < 0.001 | 0.008 | 0.03 | < 0.001 | |||||||||
0.003 | 0.016 | 0.011 | 0.000 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | ||||||||
0.001 | 0.042 | < 0.001 | 0.001 | ||||||||||||
0.029 | < 0.001 | 0.002 | 0.001 |
Holotype of
Body relatively short (
Forelimbs moderate in stature, relatively short (
Tail original, long 76.7 mm (
The specific epithet
Summary statistics from the
OTU pairs | F model | R2 | ||
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6.8474 | 0.4064 | 0.015 | 0.544 | |
8.8824 | 0.4467 | 0.003 | 0.122 | |
3.3630 | 0.2189 | 0.006 | 0.201 | |
10.4580 | 0.4874 | 0.003 | 0.114 | |
6.5983 | 0.3367 | 0.000 | 0.012 | |
6.6357 | 0.3067 | 0.000 | 0.004 | |
3.8646 | 0.1945 | 0.001 | 0.033 | |
15.4818 | 0.7559 | 0.048 | 1.000 | |
15.9186 | 0.6946 | 0.027 | 0.967 | |
19.0130 | 0.7038 | 0.022 | 0.804 | |
4.4753 | 0.4723 | 0.029 | 1.000 | |
14.9425 | 0.7135 | 0.018 | 0.643 | |
8.7953 | 0.5237 | 0.009 | 0.317 | |
14.7978 | 0.6218 | 0.006 | 0.226 | |
9.8976 | 0.6644 | 0.029 | 1.000 | |
4.5646 | 0.3947 | 0.008 | 0.278 | |
6.7120 | 0.4272 | 0.003 | 0.124 | |
5.6585 | 0.3614 | 0.002 | 0.067 | |
7.4818 | 0.5550 | 0.018 | 0.643 | |
22.8234 | 0.7405 | 0.008 | 0.283 | |
17.0146 | 0.6540 | 0.006 | 0.227 | |
17.8585 | 0.6410 | 0.001 | 0.048 | |
9.3960 | 0.4607 | 0.001 | 0.025 | |
8.2047 | 0.3869 | 0.000 | 0.005 |
Habitat of
A taxonomy consistent with evolutionary history is paramount to any downstream comparative analyses employed to address the evolution of other features of the group be they ecology, behavior, or habitat preference. By delimiting new and existing species using a phylogenetic analysis and then diagnosing those species using univariate and multivariate statistical analyses of their morphological data, phylogenetic history will not conflated with convergent evolution (Fig.
Comparison of a morphology-based taxonomy pre-2020 and the current phylogenetic-based taxonomy illustrating the misrepresentation of the diversity of the
The description of the four new species,
This work was financially supported by Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation (grant no. RGNS 64-038). NA was supported by Srinakharinwirot University Research Grant (no. 596/2564). This research was reviewed and approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (ACKU64-SCI-006) and the Department of National Parks, Wildlife and Plant Conservation, Thailand provided the research permission. We would like to thank Suchai Horradee (Chaloem Rattanakosin National Park), Charoen Jaichon (Thong Pha Phum National Park) and Mana Phermpool (Kaeng Krachan National Park) for facilitating the fieldwork. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Piyawan Puanprapai assisted with fieldwork. We would like to thank Bryan L. Stuart for suggestions on this research and Truong Nguyen and an anonymous reviewer improved the manuscript.
Data frame for the multiple factor analysis of the putative species of the
morphological data
Data frame for the multiple factor analysis of the putative species of the