Corresponding author: Anchalee Aowphol (
Academic editor: Thomas Ziegler
Yodthong S, Rujirawan A, Stuart BL, Grismer LL, Aksornneam A, Termprayoon K, Ampai N, Aowphol A (2022) A new species in the
The Bent-toed Gecko genus
The
During our fieldwork in 2019 and 2021, we collected
A total of 22 specimens (eleven adult males, nine adult females, and two juveniles) of the Si Sawat
Map illustrating the type locality at Tham Phrathat Protection Unit (yellow star) and paratype localities (yellow cycle) at Erawan Waterfall (1) and at Tham Than Lot Noi-Tham Than Lot Yai Nature Trail (2), Si Sawat District, Kanchanaburi Province, Thailand of
Total genomic DNA were extracted from preserved liver tissue of nine individuals of the Si Sawat species (Table
Specimens used in this study, including localities, museum numbers and GenBank accession numbers of the mitochondrial NADH dehydrogenase subunit 2 gene and flanking tRNA regions.
Species | Locality | Voucher No. | Accession No. | Reference |
---|---|---|---|---|
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Tha Ra Rak Waterfall, Mae Sot Dist., Tak Prov., Thailand | ZMMU R 16626 |
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|
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Khao Ramrome, Ron Phibun Dist., Nakhon Si Thammarat Prov., Thailand | AUP-00573 |
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|
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Dammathet Cave, 19.8 km east of Mawlamyine, Mawlamyine Dist., Mon State, Myanmar | LSUHC 12864 |
|
|
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Doi Suthep, Mueang Dist., Chiang Mai Prov., Thailand | AUP-00777 |
|
|
|
Chiang Dao, Chiang Mai Prov., Thailand | AUP-00768 |
|
|
|
Coral Cave, Pang Mapha Dist., Mae Hong Son Prov., Thailand | AUP-00771 |
|
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|
Nakai Dist., Khammouan Prov., Laos |
|
|
|
|
Tiger Head mountain, Doi Inthanon National Park, Chom Thong Dist., Chiang Mai Prov., Thailand | AUP-00156 |
|
|
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The proposed Lenya National Park Extension, Tanintharyi Region, Myanmar | USNM 587789 |
|
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The proposed Lenya National Park Extension, Tanintharyi Region, Myanmar | USNM 587788 |
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The proposed Lenya National Park Extension, Tanintharyi Region, Myanmar | CAS 260233 |
|
|
|
Linno Cave region 5 km south-west of Hpa-an, Hpa-an Dist., Kayin State, Myanmar | LSUHC 12825 |
|
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12.7 km north-east of Ywangan, Linn-Way Village, Yum Twing Gyi Cave, Taunggyi Dist., Shan State, Myanmar, | BYU 52214 |
|
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Tha Pha Pum Subdist., Mae La Noi Dist., Mae Hong Son Prov., Thailand |
|
|
|
|
Chaloem Rattanakosin National Park, Khao Chot Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Chaloem Rattanakosin National Park, Khao Chot Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Chaloem Rattanakosin National Park, Khao Chot Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Chaloem Rattanakosin National Park, Khao Chot Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand | – | This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
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Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Erawan National Park, Tha Kradan Subdist., Si Sawat Dist., Kanchanaburi Prov., Thailand |
|
This study | |
|
Suan Phueng Distc., Ratchburi Prov., Thailand | HLM 0307 |
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Kraburi Dist., Phang-nga Prov., Thailand | MS 460 |
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Muang Dist., Ranong Prov., Thailand | MS 585 |
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Chumpon Prov., Thailand | LSUHC 9486 |
|
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in the proposed Lenya National Park, Tanintharyi Region, Myanmar | USNM 587409 |
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in the proposed Lenya National Park, Tanintharyi Region, Myanmar | USNM 587792 |
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in the proposed Lenya National Park, Tanintharyi Region, Myanmar | USNM 587791 |
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Pharpoun Cave, 38.4 km south-east of Mawlamyine, Mawlamyine Dist., Mon State, Myanmar | BYU 52215 |
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Sadan Cave, 17 km south-east of Hpa-an, Hpa-an Dist., Kayin State, Myanmar | LSUHC 12853 |
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Sadan Sin Cave 10.5 km north-west of Mawlamyine, Mawlamyine Dist., Mon State, Myanmar | BYU 52220 |
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Sai Yok National Park, Kanchanaburi Prov., Thailand | MS 484 |
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Sai Yok National Park, Kanchanaburi Prov., Thailand | MS 480 |
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Sai Yok Dist., Kanchanaburi Prov., Thailand | AUP-00773 |
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Tham Sanook, Muang Dist., Chumphon Prov., Thailand | AUP-00570 |
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Sanpel Cave, 21.3 km south-east of Mawlamyine, Mawlamyine Dist., Mon State, Myanmar | LSUHC 12886 |
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5.3 km north of Pyinyaung Village at the Apache Cement factory mining site, Mandalay Region, Myanmar | BYU 52227 |
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Moe Cham Pae Dist., Mae Hong Son Prov., Thailand | HLM 0357 |
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Krabi, Trang Prov., Thailand | HLM 0358 |
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Thong Pha Phum National Park, Thong Pha Phum Dist., Kanchanaburi Prov., Thailand | AUP-01718 |
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Near Vajiralongkorn dam, Thong Pha Phum National Park, Thong Pha Phum Dist., Kanchanaburi Prov., Thailand | AUP-01722 |
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Tham Khao Sonk hill, Surat Thani Prov., Thailand |
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Tham Khao Sonk hill, Surat Thani Prov., Thailand |
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Ban Tha Sao, Sai-Yok Dist., Kanchanaburi Prov., Thailand | IRSNB2380 |
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Wang Krachae Subdist., Sai Yok Dist., Kanchanaburi Prov., Thailand | AUP-00776 |
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Yathe Pyan Cave, 9 km south-west of Hpa-an, Hpa-an Dist., Kayin State, Myanmar | LSUHC 12823 |
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Mueang Krabi, Krabi, Thailand | HLM 0344 |
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Khao Luang National Park, Thailand | CUMZR 2005.07.30.54 |
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Ron Phibun, Nakhon Si Thammarat, Thailand | – | This study | |
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||||
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Thong Pha Phum National Park, Thong Pha Phum Dist., Kanchanaburi Prov., Thailand | AUP-01724 |
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Shwesettaw wildlife sanctuary, Mimbu Township, Magway Div., Myanmar | CAS 213628 |
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Thong Pha Phum National Park, Thong Pha Phum Dist., Kanchanaburi Prov., Thailand | AUP-01710 |
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Rathegala, Sri Lanka | AMB 7420 |
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|
All available
Models of molecular evolution selected for the maximum likelihood and Bayesian analyses.
Gene | Model selected | Model applied for |
Model applied for |
---|---|---|---|
|
|||
1st position | TVM+F+I+G4 | TVM+F+I+G4 | GTR+I+Γ |
2nd position | TPM3u+F+I+G4 | TPM3u+F+I+G4 | GTR+I+Γ |
3rd position | TIM3+F+G4 | TIM3+F+G4 | GTR+I+Γ |
|
TIMe+G4 | TIMe+G4 | GTR+I+Γ |
The
Mensural, meristic, and qualitative characters were taken using a Nikon SMZ 745 Zoom Stereomicroscope. Measurements were taken on the left side of the body when possible, with digital calipers (Mitutoyo CD-6” ASX Digimatic Caliper, Japan) to the nearest 0.1 mm. Characters and abbreviations were modified from
Meristic characters were taken on both right and left (R/L) sides when possible. The presence, absence, and/or numbers of the characters were recorded as follows:
Non-meristic morphological characters examined were the degree of body tuberculation, weak tuberculation refers to low and weakly keeled dorsal body tubercles whereas prominent tuberculation refers to raised and prominently keeled dorsal body tubercles; body tubercles extending past the base of the tail or not; enlarged femoral scales and precloacal scales contiguous or separated by a diastema at the base of the femora; a precloacal depression or groove present or absent; transversely expanded, median subcaudal scales present or absent; and the relative length to width ratio of the transversely expanded, median subcaudal scales. Color pattern characters evaluated were the nuchal loop being continuous from eye to eye, or separated medially into paravertebral blotches; the dorsal body bands bearing paired, paravertebral elements or fused medially; dark dorsal body bands edged with light-colored tubercles or not; dark markings present or absent in the dorsal interspace; ventrolateral body folds weak or prominent; top of head bearing combinations of dark diffuse mottling or dark distinct blotches overlain with a light-colored reticulating network or not; light-colored caudal bands encircling tail or not; and regenerated tail bearing a pattern of distinct, dark spots or not.
Morphological comparisons were based on examination of the holotype of
The total aligned dataset contained 1,444 mtDNA characters with gaps from 50 individuals of
The best maximum likelihood tree showing the relationships of the
In both analyses, the Si Sawat species represented a deeply divergent mtDNA lineage and a strongly supported monophyletic group (1.0
The uncorrected pairwise sequence divergences (
Uncorrected pairwise sequence divergences (
No. | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
|
0.3 | |||||||||||||
( |
(0.0–1.2) | |||||||||||||
|
7.7 | – | ||||||||||||
( |
(7.7–8.0) | |||||||||||||
|
11.5 | 10.8 | – | |||||||||||
( |
(11.2–11.7) | (10.8) | ||||||||||||
|
11.6 | 11.2 | 1.8 | – | ||||||||||
( |
(11.4–11.7) | (11.2) | (1.8) | |||||||||||
|
11.8 | 10.0 | 9.3 | 9.3 | – | |||||||||
( |
(11.7–12.0) | (10.0) | (9.3) | (9.3) | ||||||||||
|
12.0 | 11.3 | 9.0 | 9.2 | 9.7 | 0.0 | ||||||||
( |
(11.4–12.4) | (11.1–11.4) | (9.0) | (9.1–9.2) | (9.7) | (0.0) | ||||||||
|
12.3 | 11.1 | 10.6 | 10.9 | 7.4 | 8.3 | 0.1 | |||||||
( |
(11.7–13.2) | (10.7–11.9) | (10.2–11.4) | (10.5–11.6) | (7.1–8.1) | (7.7–9.7) | (0.0–0.1) | |||||||
|
12.7 | 12.1 | 9.9 | 9.6 | 6.0 | 7.1 | 8.02 | – | ||||||
( |
(12.6–12.8) | (12.1) | (9.9) | (9.6) | (6.0) | (7.0–7.2) | (7.7–8.7) | |||||||
|
12.8 | 11.3 | 9.7 | 9.9 | 6.5 | 6.9 | 7.7 | 7.7 | – | |||||
( |
(12.7–12.9) | (11.3) | (9.7) | (9.9) | (6.5) | (6.6–7.2) | (7.4–8.3) | (7.7) | ||||||
|
12.3 | 11.6 | 9.7 | 9.53 | 5.49 | 7.17 | 9.1 | 7.4 | 7.8 | – | ||||
( |
(12.13–12.57) | (11.58) | (9.7) | (9.53) | (5.49) | (6.61–7.18) | (8.63–9.97) | (7.43) | (7.8) | |||||
|
13.0 | 12.1 | 9.3 | 9.3 | 10.7 | 10.8 | 11.2 | 11.0 | 10.6 | 11.4 | – | |||
( |
(12.7–13.2) | (12.1) | (9.3) | (9.3) | (10.7) | (10.4–11.2) | (10.9–11.7) | (11.0) | (10.6) | (11.4) | ||||
|
15.4 | 14.6 | 14.2 | 14.8 | 14.4 | 14.9 | 14.8 | 15.1 | 14.6 | 15.5 | 15.3 | 0.1 | ||
( |
(15.2–15.6) | (14.3 –14.8) | (14.1–14.3) | (14.7–14.9) | (14.2–14.6) | (14.6–15.2) | (14.1–16.4) | (14.9–15.3) | (14.4–14.8) | (15.2–15.8) | (15.0–15.5) | (0.1–0.2) | ||
|
16. | 15.1 | 16.5 | 16.6 | 15.5 | 14.9 | 16.3 | 16.9 | 16.1 | 16.7 | 17.5 | 16.1 | 0.5 | |
( |
(16.6–17.0) | (15.0–15.1) | (16.4–16.6) | (16.4–16.7) | (15.4–15.7) | (14.1–15.6) | (15.6–17.6) | (16.7–17.1) | (16.0–16.1) | (16.6–16.8) | (17.3–17.6) | (15.7–16.4) | (0.2–0.6) | |
|
17.6 | 16.3 | 16.8 | 17.0 | 16.9 | 16.3 | 17.7 | 17.5 | 17.3 | 17.2 | 17.4 | 17.2 | 13.2 | 6.0 |
( |
(17.3–17.7) | (16.2–16.3) | (16.6–16.9) | (16.9–17.0) | (16.8–16.9) | (15.3–17.2) | (16.7–19.2) | (17.5) | (17.2–17.4) | (17.2–17.4) | (17.2–17.6) | (15.7–16.4) | (13.0–13.2) | (6.0) |
Based on the results of mtDNA and morphological comparisons (see below), the
Adult male holotype of
Adult male holotype of
Paratypes of
One juvenile (
The specific epithet
(Figs
Body slender, relatively short (
Limbs moderately slender; forelimbs relatively short (
Tail 58.1 mm in length, original, slightly longer than
Morphometric, meristic and color pattern characters of the type series and referred specimens of
Descriptive measurements in millimeters of the type series of
Characters | Holotype male | Holotype and paratype males | Paratype females | ||
---|---|---|---|---|---|
Min–Max | Mean ± SD | Min–Max | Mean ± SD | ||
|
56.4 | 53.7–63.3 | 58.0 ± 3.4 | 58.6–75.8 | 68.7 ± 5.6 |
|
16.4 | 15.5–18.10 | 16.6 ± 0.9 | 16.8–22.0 | 19.3 ± 1.7 |
|
10.6 | 10.1–12.1 | 11.0 ± 0.6 | 11.5–15.4 | 13.3 ± 1.2 |
|
6.5 | 5.8–7.5 | 6.6 ± 0.6 | 6.4–9.1 | 7.7 ± 0.9 |
|
4.8 | 4.3–5.3 | 4.7 ± 0.4 | 4.8–5.4 | 5.2 ± 0.3 |
|
5.1 | 4.0–5.3 | 4.9 ± 0.4 | 5.1–6.7 | 5.9 ± 0.6 |
|
6.6 | 5.9–7.4 | 6.6 ± 0.4 | 6.5–8.8 | 7.4 ± 0.7 |
|
5.2 | 4.3–5.4 | 4.9 ± 0.3 | 4.8–6.5 | 5.5 ± 0.5 |
|
5.3 | 5.1–6.4 | 5.7 ± 0.4 | 5.7–7.6 | 6.6 ± 0.7 |
|
1.5 | 1.1–1.8 | 1.4 ± 0.2 | 1.3–1.7 | 1.5 ± 0.2 |
|
1.9 | 1.7–2.1 | 1.9 ± 0.1 | 1.8–2.4 | 2.1 ± 0.2 |
|
24.7 | 22.3–27.8 | 25.4 ± 1.7 | 26.1–34.4 | 31.1 ± 2.8 |
|
8.3 | 8.0–9.6 | 8.8 ± 0.5 | 9.1–11.8 | 10.4 ± 1.0 |
|
10.4 | 9.8–11.7 | 10.8 ± 0.7 | 11.4–14.1 | 12.8 ± 1.0 |
|
58.1 | 58.1–62.10a | 59.9 ± 1.7a | 64.0–77.7c | 71.4 ± 5.7c |
|
NA | 41.8–60.0b | 49.5 ± 5.8b | 25.7–55.1d | 42.8 ± 12.9d |
|
4.6 | 4.0–5.4 | 4.9 ± 0.5 | 4.5–5.3 | 4.9 ± 0.3 |
|
4.0 | 4.0–5.6 | 4.7 ± 0.5 | 4.3–5.4 | 4.8 ± 0.4 |
a
Meristic characters (right/left) and color patterns of
Characters | Holotype | Holotype and paratypes |
---|---|---|
Min–Max | ||
|
12/11 | 10–13 |
|
8/7 | 6–9 |
|
9/10 | 8–11 |
|
5/7 | 5–8 |
Body tubercles pointed and keeled | Yes | Yes |
|
28 | 25–34 |
|
16 | 16–21 |
|
38 | 34–42 |
|
12/12 | 9–12 |
|
4/4 | 3–5 |
|
16/16 | 12–16 |
|
13/13 | 10–13 |
|
5/6 | 4–6 |
|
18/19 | 15–19 |
Enlarge femoral and precloacal scales continuous | Yes | Yes |
|
33 | 30–39* |
|
Absent | Absent |
|
4 | 4–5 |
|
2/2 | 2–3 |
Enlarged median subcaudal scales | No | No |
Nuchal loop discontinuous | Yes | Yes & No |
Paravertebral elements not in contact | Yes | Yes |
|
6 | 4–7 |
|
11 | 9–12 |
|
11 | 9–12 |
*
Meristic characters (right/left) and color patterns of the referred specimens of
Characters | Min–Max | ||
---|---|---|---|
Age | Juvenile | Juvenile | |
|
40.6 | 31.3 | 31.3–40.6 |
|
12/11 | 12/10 | 10–12 |
|
8/7 | 8/7 | 7–8 |
|
10/9 | 8/8 | 8–10 |
|
7/6 | 5/6 | 5–7 |
Body tubercles pointed and keeled | Yes | Yes | Yes |
|
22 | 27 | 22–27 |
|
16 | 17 | 16–17 |
|
34 | 40 | 34–40 |
|
11/11 | 10/10 | 10–11 |
|
4/4 | 4/4 | 4 |
|
15/15 | 14/14 | 14–15 |
|
12/12 | 11/11 | 11–12 |
|
6/5 | 6/6 | 5–6 |
|
18/17 | 17/17 | 17–18 |
Enlarge femoral and precloacal scales continuous | Yes | Yes | Yes |
|
33 | 33 | 33 |
|
5 | 5 | 5 |
|
2/3 | 2/2 | 2–3 |
Enlarged median subcaudal scales | NA | No | No |
Nuchal loop discontinuous | Yes | Yes | Yes |
Paravertebral elements not in contact | Yes | Yes | Yes |
|
6 | 5 | 5–6 |
|
NA | 10 | 10 |
|
NA | 11 | 11 |
Internasal scales of ten specimens (
Sampling localities of
At Tham Phrathat Protection Unit, the holotype (
Habitat of
At Erawan Waterfall, one gravid female (
At Tham Than Lot Noi-Tham Than Lot Yai Nature Trail, one juvenile specimen (
Adult male holotype of
The combination of morphological and molecular phylogenetic evidence in this study corroborated the hypothesis that the Si Sawat population should be recognized as a distinct species, described here as
This work was financially supported by the Office of the Ministry of Higher Education, Science, Research and Innovation; and the Thailand Science Research and Innovation through the Kasetsart University Reinventing University Program 2021. AR and AA were supported by Kasetsart University Research and Development Institute (KURDI), the Department of Zoology, Faculty of Science, Kasetsart University and 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). The study was ethical reviews and approvals by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project number ACKU61-SCI-008 and ACKU64-SCI-005) and the Department of National Parks, Wildlife and Plant Conservation, Thailand provided research permission (No. 0907.4/3328 and No. 0907.4/20535). We would like to thank Suchai Horradee (Chaloem Rattanakosin National Park), Porayoot Vaivong and Peerawat Sirothphiphat (Erawan National Park) for facilitating the fieldwork. Sang Nguyen and Han Ngo improved the manuscript.