Corresponding authors: Vinh Quang Luu (
Academic editor: Thomas Ziegler
Integrated analyses using maximum likelihood (
Luu VQ, Nguyen TH, Le MD, Grismer JL, Ha HB, Sitthivong S, Hoang TT, Grismer LL (2023) Two new species of
The genus
Location of the type localities of all known species of
During a recent herpetofaunal surveys in Chu Se Mountain Pass, Hbong Commune, Gia Lai Province in Vietnam and Vientiane Province in Laos, new populations of
A total of six
The general lineage concept (
Four samples of the newly collected specimens were analyzed, two from Gia Lai Province, Vietnam (
We obtained 1,444 base pairs of NADH dehydrogenase subunit 2 gene (
List of specimens used for the phylogenetic analyses.
Species | Catalog no. | Location | GenBank no. |
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ZFMK87274 | Nui Chua NP, Ninh Thuan Province, southern Vietnam |
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Chu Se District, Gia Lai Province, Vietnam |
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Chu Se District, Gia Lai Province, Vietnam |
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Khammouane Province, Laos |
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IEBR A.2019.5 | Khammouane Province, Laos |
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IEBR A.2019.6 | Khammouane Province, Laos |
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VU 022 | Captive, Thailand |
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ZFMK 97745 | Vinh Cuu, Dong Nai Province, Vietnam |
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Muangfuang District, Vientiane Province, Central Laos |
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Muangfuang District, Vientiane Province, Central Laos |
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VU 023 | Captive, Thailand |
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LSUHC 7328 | Phnom Aural, Purset Province, Cambodia |
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FMNH 263003 | Keo Seima District, Mondolkiri- Province, Cambodia |
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LSUHC 7378 | Phnom Aural, Purset Province, Cambodia |
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Nasaithong District, Vientiane Capital, Laos |
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Nasaithong District, Vientiane Capital, Laos |
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Nasaithong District, Vientiane Capital, Laos |
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Vientiane Capital, Laos |
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Vientiane Capital, Laos |
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Vientiane Capital, Laos |
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Vientiane Capital, Laos |
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LSUHC 9466 | Sai Yok, Kanchanaburi Province, Thailand |
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ZFMK 96680 | Phu Quy Island, Binh Thuan Province, Vietnam |
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CAS 257300 | Phu Quy Island, Binh Thuan Province, Vietnam |
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IEBR A 2014-26 | Phu Quy Island, Binh Thuan Province, Vietnam |
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IEBR A 2014-27 | Phu Quy Island, Binh Thuan Province, Vietnam |
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ZFMK 87273 | Nui Chua, Ninh Thuan Province, Vietnam |
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IEBR R.20163 | Nha Trang, Khánh Hòa Province, Vietnam |
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Maximum likelihood (
A time-calibrated Bayesian phylogenetic tree was estimated using
The morphological data set comprised six closely related species including six type specimens of
Morphological data included both meristic and morphometric characters. Morphological characters were taken from the 44 specimens following
Meristic data taken were:
Color pattern on dorsum including the presence or absence of canthal stripes (
Sex and raw meristic and categorical data used in the analyses from specimens of
Species | Museum no. | Sex | Meristic data | Categorical data | |||||||||
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V | Canthal stripe | Lips strong barred | Blotches on the head round | Blotches on dorsum round | Two regularly disposed whitish tubercles on each side of the flanks | ||||||
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IEBR A.0802 | m | 8 | 6 | 6 | 10 | 22 | 14 | present | no | yes | yes | absent |
ZFMK 97746 | m | 8 | 6.5 | 6 | 10 | 23 | 14.5 | present | no | yes | yes | absent | |
IEBR A.0801 | f | 8.5 | 7 | 6 | 10 | 22 | 12 | present | no | yes | yes | absent | |
ZFMK 97745 | f | 7.5 | 6 | 5.5 | 10 | 23 | 13 | present | no | yes | yes | absent | |
VNMN R.2016.1 | f | 8 | 6 | 5.5 | 8 | 23 | 15 | present | no | yes | yes | absent | |
VNMN R.2016.2 | f | 8 | 6.5 | 6 | 7 | 20 | 13 | present | no | yes | yes | absent | |
m | 7.5 | 6 | 6 | 7 | 21 | 14 | present | yes | yes | yes | present | ||
f | 7 | 6 | 6 | 7 | 19 | 14 | present | yes | yes | yes | present | ||
mj | 8 | 7 | 6 | 7 | 21 | 14.5 | present | yes | yes | yes | present | ||
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ZRC 2.6024 | m | 5 | 6 | 5 | 10 | 20 | 13 | present | no | no | no | present |
ZRC 2.6025 | m | 5 | 6 | 5 | 9 | 20 | 13 | present | no | no | no | present | |
ZRC 2.6026 | j | 5 | 6 | 6 | 8 | 20 | 13 | present | no | no | no | present | |
ZRC 2.6027 | j | 6 | 7 | 6 | 8 | 20 | 13 | present | no | no | no | present | |
IEBR R.2016.3 | m | 8 | 6 | 5.5 | 10 | 19 | 13.5 | present | no | no | no | present | |
VNMN R.2016.3 | m | 7.5 | 6 | 5.5 | 9 | 19 | 13.5 | present | no | no | no | present | |
IEBR R.2016.1 | f | 7 | 6 | 5.5 | 8 | 18 | 13.5 | present | no | no | no | present | |
VNMN R.2016.4 | f | 7.5 | 7 | 6 | 9 | 20 | 13 | present | no | no | no | present | |
ZFMK 97748 | f | 7.5 | 6 | 6 | 8 | 20 | 14 | present | no | no | no | present | |
ZFMK 97747 | mj | 7.5 | 6 | 5.5 | 10 | 15 | 13.5 | present | no | no | no | present | |
IEBR R.2016.4 | f j | 8 | 7 | 6 | 7 | 21 | 12.5 | present | no | no | no | present | |
ZFMK 97749 | fj | 7 | 6.5 | 5.5 | 8 | 19 | 13.5 | present | no | no | no | present | |
sp. | m | 8 | 6.5 | 6 | 9 | 24 | 14 | present | no | no | yes | present | |
f | 7.5 | 5.5 | 6 | 8 | 23 | 14.5 | present | no | no | yes | present | ||
fj | 8 | 7 | 6 | 8 | 23 | 14 | present | no | no | yes | present | ||
f j | 8.5 | 6 | 6 | 8 | 22 | 13.5 | present | no | no | yes | present | ||
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m | 7 | 5 | 6 | 8 | 24 | 14 | present | yes | no | no | present | |
m | 8 | 6 | 6 | 8 | 23 | 15 | present | yes | no | no | present | ||
m | 8 | 5.5 | 6 | 8 | 23 | 15 | present | yes | no | no | present | ||
f | 8 | 5.5 | 6 | 8 | 23 | 15 | present | yes | no | no | present | ||
f | 8 | 6 | 6 | 7 | 26 | 13 | present | yes | no | no | present | ||
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LSUHC09284 | f | 8 | 7 | 6 | 9 | 19 | 14 | absent | yes | no | yes | present |
LSUHC08522 | f | 8 | 6.5 | 6 | 10 | 22 | 14.5 | absent | yes | no | yes | present | |
LSUHC08487 | f | 8 | 7 | 6 | 10 | 20 | 14.5 | absent | yes | no | yes | present | |
LSUHC08420 | m | 8.5 | 7 | 6 | 10 | 21 | 13 | absent | yes | no | yes | present | |
LSUHC08491 | f | 8 | 7 | 6 | 9 | 20 | 14.5 | absent | yes | no | yes | present | |
LSUHC07328 | j | 7.5 | 6 | 5.5 | 9 | 22 | 14 | absent | yes | no | yes | present | |
LSUHC07378 | m | 8 | 6 | 6 | 10 | 20 | 14.5 | absent | yes | no | yes | present | |
LSUHC09289 | m | 7.5 | 6 | 6 | 10 | 21 | 16 | absent | yes | no | yes | present | |
m | 7 | 6.5 | 6 | 7 | 21 | 15 | absent | yes | no | no | present | ||
m | 8 | 7 | 6 | 7 | 20 | 15 | absent | yes | no | no | present | ||
f | 8 | 6.5 | 6 | 7 | 21 | 15 | absent | yes | no | no | present | ||
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m | 9.5 | 8 | 7.5 | 9 | 23 | 15 | absent | yes | no | no | absent | |
IEBR A.2019.5 | f | 8.5 | 8 | 7 | 8 | 23 | 15 | absent | yes | no | no | absent | |
IEBR A.2019.6 | f | 9 | 7.5 | 8 | 8 | 24 | 15 | absent | yes | no | no | absent |
Sex and raw morphometric data used in the analyses from specimens of
Species | Museum no. | Sex |
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FAr | TBLr | AGr |
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IEBR A.0802 | m | 43.9 | 9.4 | 7.3 | 7.7 | 4.7 | 1.5 | 2.7 | 3.7 | 5 | 3.5 | 1.6 | 4 | 6.2 | 7.7 | 18.7 |
ZFMK 97746 | m | 40.6 | 8.5 | 6.7 | 6 | 4.3 | 1.3 | 2.2 | 3.2 | 4.4 | 3.6 | 1.3 | 3.5 | 6.7 | 7 | 18.2 | |
IEBR A.0801 | f | 45.9 | 9.7 | 7.2 | 6.6 | 5.2 | 1.2 | 3.3 | 3.4 | 4.4 | 3.4 | 1.5 | 3.7 | 5.9 | 7.2 | 21.2 | |
ZFMK 97745 | f | 47.5 | 9.6 | 7.6 | 6.8 | 4.7 | 1.5 | 3.1 | 3.5 | 4.9 | 3.9 | 1.5 | 3.7 | 6 | 7.3 | 21.5 | |
VNMN R.2016.1 | f | 43.3 | 9.3 | 7.1 | 6.5 | 4.4 | 1.3 | 2.5 | 3.5 | 4.6 | 3.8 | 1.5 | 3.8 | 6.1 | 7.5 | 20.6 | |
VNMN R.2016.2 | f | 46.7 | 9.2 | 7.7 | 6.2 | 4.6 | 1.2 | 3.1 | 3.8 | 5.2 | 3.6 | 1.5 | 3.4 | 6.6 | 7 | 30.3 | |
m | 41.2 | 8.6 | 11.7 | 7.7 | 5.2 | 1.1 | 2.9 | 3.1 | 4.3 | 3.3 | 1.3 | 1.2 | 6.1 | 6.9 | 15.8 | ||
f | 47.4 | 8.4 | 12.3 | 8.8 | 6.1 | 1.3 | 3.3 | 3.5 | 4.8 | 3.5 | 1.5 | 1.4 | 6.3 | 7.7 | 21.8 | ||
mj | 35.9 | 8.3 | 10.9 | 6.8 | 4.7 | 0.9 | 2.6 | 2.9 | 3.8 | 3 | 1.3 | 1.3 | 4.5 | 5.6 | 14 | ||
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ZRC 2.6024 | m | 40.8 | 8 | 7.5 | 7.9 | 5.5 | 1 | 2.9 | 3.2 | 4.3 | 3.8 | 2.1 | 3.6 | 5.6 | 7.7 | 21 |
ZRC 2.6025 | m | 42.4 | 9.1 | 7.5 | 7.6 | 6 | 1.1 | 2.9 | 3.7 | 4.6 | 4 | 1.6 | 3.6 | 6.2 | 7.2 | 21 | |
ZRC 2.6026 | j | 26.6 | 5.4 | 5.4 | 5.2 | 4 | 0.6 | 2.1 | 2.2 | 3 | 2.5 | 1.1 | 2.7 | 4.4 | 4.4 | 13 | |
ZRC 2.6027 | j | 25.9 | 4 | 5.2 | 5.1 | 3.3 | 0.6 | 1.8 | 2.3 | 3.5 | 2.2 | 0.9 | 2.1 | 4 | 4.6 | 11.8 | |
IEBR R.2016.3 | m | 39 | 6.5 | 6.9 | 6.8 | 4.2 | 1.1 | 2.8 | 2.9 | 3.9 | 3.1 | 1.1 | 1.6 | 4.7 | 6.5 | 14.8 | |
VNMN R.2016.3 | m | 39.9 | 7.8 | 7.2 | 7 | 4.7 | 0.8 | 2.5 | 3.4 | 4.6 | 3.3 | 1.3 | 1.7 | 5.2 | 6.5 | 16.6 | |
IEBR R.2016.1 | f | 43.5 | 7.6 | 7.6 | 6.9 | 4.7 | 1 | 2.7 | 3.1 | 4.5 | 2.7 | 1.3 | 1.6 | 5 | 6.2 | 19.2 | |
VNMN R.2016.4 | f | 43.7 | 8.6 | 7.7 | 7.7 | 4.7 | 1.1 | 2.8 | 3.4 | 4.7 | 3.6 | 1.3 | 1.8 | 5.5 | 6.3 | 18.2 | |
ZFMK 97748 | f | 45.2 | 10.3 | 8.5 | 8.2 | 5.7 | 1.3 | 2.9 | 3.8 | 5.4 | 4.3 | 1.5 | 2.7 | 5.5 | 6.6 | 19.2 | |
ZFMK 97747 | mj | 34.1 | 4.7 | 6.2 | 5.7 | 4.1 | 0.9 | 2.3 | 2.7 | 3.6 | 2.5 | 1.2 | 1.3 | 4.2 | 5.9 | 12.3 | |
IEBR R.2016.4 | f j | 31.2 | 5.2 | 5.7 | 5.8 | 3.7 | 0.9 | 2.4 | 2.4 | 3.4 | 2.4 | 1 | 1.2 | 3.1 | 4.8 | 11.9 | |
ZFMK 97749 | fj | 29.2 | 4.8 | 5.2 | 4.8 | 3.1 | 0.9 | 2.5 | 2.1 | 2.9 | 2.3 | 1 | 1.2 | 3.1 | 4.9 | 11.1 | |
sp. | m | 46 | 10.7 | 13.7 | 7.8 | 5.7 | 1.4 | 2.2 | 3.4 | 4.8 | 3.7 | 1.7 | 1.9 | 5.5 | 6.8 | 20.2 | |
f | 35.2 | 7.5 | 10.9 | 6.9 | 3.3 | 1.1 | 2.1 | 2.9 | 4.1 | 3.2 | 1.3 | 1.6 | 4.7 | 6.2 | 14.2 | ||
fj | 31.1 | 5.1 | 9.2 | 5.1 | 2.4 | 1.2 | 2.2 | 2.3 | 2.3 | 2.5 | 1.1 | 1.4 | 4.5 | 5.1 | 12.5 | ||
f j | 30.3 | 5.6 | 9.3 | 5.8 | 3 | 0.8 | 2 | 2.7 | 3.4 | 2.9 | 1.1 | 1.5 | 3.7 | 4.6 | 13.3 | ||
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m | 43.8 | 9.4 | 12.2 | 8.5 | 5.6 | 1.6 | 3.4 | 3 | 5.1 | 3.4 | 1.3 | 1.7 | 6.2 | 7.3 | 20.5 | |
m | 47.1 | 11.1 | 12.9 | 9.7 | 5.9 | 1.9 | 3.3 | 3.4 | 5 | 3.5 | 1.6 | 1.8 | 5.6 | 8 | 19.5 | ||
m | 39.8 | 8.9 | 11.6 | 7.9 | 5.2 | 1.2 | 2.9 | 2.9 | 4.2 | 3.1 | 1.7 | 1.3 | 4.8 | 6.6 | 17.4 | ||
f | 35.5 | 9.4 | 9.7 | 6.9 | 4.2 | 1.2 | 2.2 | 2.8 | 3.7 | 2.5 | 1.2 | 1.4 | 4.3 | 5.5 | 15.5 | ||
f | 39.9 | 8.9 | 11.4 | 7.6 | 4.4 | 1.5 | 3.1 | 2.6 | 4 | 3.1 | 1.5 | 1.3 | 4.9 | 6 | 19.7 | ||
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LSUHC09284 | f | 45.4 | 8.6 | 12.8 | 8.7 | 5.2 | 1.6 | 3 | 3.7 | 5.1 | 4 | 2 | 3.6 | 6.7 | 7.3 | 19 |
LSUHC08522 | f | 44.1 | 9.4 | 12.5 | 8.1 | 5.7 | 1.4 | 2.4 | 4.4 | 5.2 | 4.5 | 1.8 | 3.7 | 6.7 | 6.9 | 21.2 | |
LSUHC08487 | f | 48.6 | 10.7 | 14.3 | 8.7 | 5.4 | 1.6 | 3.2 | 3.4 | 5.4 | 4.2 | 1.7 | 3.5 | 7.1 | 8 | 21.8 | |
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LSUHC08420 | m | 46.9 | 8.8 | 13.1 | 9.1 | 5.3 | 1.3 | 2.7 | 3.7 | 5.3 | 3.9 | 1.5 | 3.7 | 6.7 | 7.3 | 20.7 |
LSUHC08491 | f | 45.2 | 10.2 | 13 | 8.2 | 5.7 | 1.4 | 2.8 | 3.3 | 4.7 | 4.2 | 2 | 3.7 | 6.2 | 6.9 | 19 | |
LSUHC07328 | j | 28.6 | 5.8 | 8.4 | 5.5 | 3 | 0.7 | 2.1 | 2.4 | 3.3 | 2.8 | 1.5 | 2.8 | 3.8 | 5 | 12 | |
LSUHC07378 | m | 36.7 | 6.5 | 10.9 | 7.3 | 4.5 | 1.3 | 2.6 | 3.1 | 4.6 | 3.4 | 1.6 | 3.4 | 6 | 6.6 | 16.1 | |
LSUHC09289 | m | 45.3 | 9.1 | 12.7 | 8.6 | 5.1 | 1.6 | 2.6 | 3.7 | 5 | 3.6 | 2 | 3.5 | 7 | 7.3 | 18.9 | |
m | 38.3 | 7.83 | 10.5 | 7.2 | 4.3 | 0.8 | 2.4 | 2.8 | 3 | 3.4 | 1.3 | 1.7 | 4.3 | 4.9 | 16.5 | ||
m | 55.6 | 11.93 | 15.2 | 10.8 | 6.9 | 2.3 | 3 | 3.8 | 5.9 | 5.1 | 1.6 | 2.3 | 6.8 | 7.2 | 23.1 | ||
f | 56.7 | 12.23 | 16.7 | 10.7 | 6.9 | 2.1 | 3.5 | 3.8 | 5.8 | 5.1 | 1.7 | 2.4 | 7.1 | 7.3 | 27.4 | ||
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m | 50.1 | 9.7 | 14.1 | 9.2 | 5.3 | 1.4 | 3.6 | 4.4 | 5.6 | 4.1 | 1.7 | 1.7 | 6.9 | 7.6 | 20.6 | |
IEBR A.2019.5 | f | 55.4 | 11.5 | 14.3 | 9.7 | 6.2 | 1.7 | 3.6 | 4 | 5.5 | 4.4 | 1.8 | 1.5 | 7.1 | 8.5 | 22.2 | |
IEBR A.2019.6 | f | 35.8 | 7.2 | 9.9 | 7 | 4 | 1.1 | 2.7 | 2.8 | 3.6 | 2.6 | 1.1 | 1.1 | 4.6 | 5.9 | 15.2 |
All statistical analyses were performed using R v. 4.2.1 (R Core Team, 2021). Morphometric characters used in the statistical analyses were
A Levene’s test for normalized morphometric and meristic characters was conducted to test for equal variances across all groups. Analyses of variance (
Morphospatial positions were visualized using principal component analysis (
To test and further corroborate the
The results of
Mean percentages of uncorrected pairwise sequence divergence (p-distances) among the species of
cf. |
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cf. |
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cf. |
6.33 |
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18.52 | 18.37 |
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11.49 | 13.16 | 16.07 |
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12.12 | 13.67 | 18.84 | 6.58. |
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cf. |
12.12 | 12.43 | 18.31 | 7.36 | 2.57 |
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10.78 | 8.17 | 18.17 | 11.36 | 12.79 | 12.50 |
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8.46 | 9.26 | 16.66 | 10.90 | 11.90 | 11.39 | 3.10 |
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13.97 | 15.33 | 17.56. | 13.35 | 14.13 | 13.92 | 13.23 | 13.24 |
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13.51 | 14.27 | 15.73 | 11.78 | 13.40 | 13.18 | 13.11 | 10.90 | 3.60 |
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13.71 | 14.83 | 16.14 | 11.74 | 12.33 | 12.22 | 12.70 | 11.96 | 12.54 | 10.56 |
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13.31 | 12.66 | 17.73 | 12.90 | 12.57 | 12.40 | 12.40 | 12.27 | 12.24 | 10.63 | 9.07 |
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13.30 | 13.04 | 15.16 | 11.23 | 13.12 | 13.01 | 11.99 | 10.70 | 14.09 | 11.53 | 12.10 | 11.09 |
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Maximum likelihood topology of the
The time-calibrated
The first two principal components (PC1 and PC2) of the
Summary statistics of the principal component analysis of
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9.060294267 | 2.842183612 | 1.53635844 | 1.413798721 | 1.291219096 | 0.903747223 | 0.850826887 |
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-0.183137642 | 0.011423135 | -0.069418522 | 0.076025214 | -0.119546371 | 0.451176774 | -0.589642945 |
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-0.287276767 | 0.064974951 | 0.187163981 | -0.199453201 | 0.019777911 | -0.133068566 | -0.114357041 |
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-0.222534372 | 0.251387029 | 0.23514022 | 0.300194841 | 0.119329084 | -0.056134295 | 0.150350725 |
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-0.264923454 | 0.100856053 | 0.274888978 | 0.257153297 | -0.193697896 | -0.025433828 | 0.20364848 |
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-0.239223187 | -0.126312635 | 0.224210506 | -0.024761051 | -0.413575793 | 0.029259717 | 0.233903564 |
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-0.2480955 | 0.169750915 | 0.178082873 | 0.002208353 | 0.210266124 | 0.137233441 | -0.232577889 |
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-0.202876478 | 0.122593123 | 0.079950567 | -0.239727042 | -0.47584928 | 0.235373894 | -0.048044975 |
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-0.265593548 | -0.130857091 | -0.293077474 | 0.03772842 | 0.029146276 | -0.105240363 | 0.142115916 |
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-0.267303156 | -0.128737264 | -0.102433066 | -0.036514974 | -0.068578553 | -0.009743264 | 0.195931987 |
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-0.276238196 | -0.150072094 | -0.016576264 | 0.149081788 | 0.006956725 | -0.271219422 | -0.084638951 |
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-0.239210846 | -0.181935095 | 0.070874242 | 0.114696597 | 0.170327297 | 0.022503069 | 0.138746934 |
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-0.131045671 | -0.460675273 | -0.164479294 | -0.032535496 | 0.242758327 | -0.169473493 | -0.152061581 |
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-0.279019143 | -0.171574811 | -0.122828868 | -0.090763378 | 0.096076353 | -0.023168457 | 0.04857391 |
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-0.256167278 | -0.099347048 | -0.096744886 | -0.219547386 | 0.043230096 | 0.332696024 | 0.101539921 |
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-0.262180808 | -0.1304743 | 0.000207287 | -0.261650023 | -0.044763987 | -0.216118201 | -0.26353247 |
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-0.138456955 | 0.383331303 | -0.225322477 | 0.206591458 | 0.113507526 | 0.110869199 | -0.176892801 |
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-0.089464182 | 0.168661032 | -0.585083828 | 0.180041929 | -0.237863864 | -0.199925005 | -0.127174583 |
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-0.156905954 | 0.393579439 | -0.305813247 | 0.010226561 | -0.089299256 | -0.164453012 | 0.254918394 |
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-0.068091843 | -0.230600144 | 0.078301763 | 0.673617186 | 0.047911701 | 0.164434462 | -0.062199104 |
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-0.140473075 | 0.310134826 | 0.27924011 | -0.11493361 | 0.255279707 | -0.405751544 | -0.23957707 |
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-0.152382721 | 0.157971329 | -0.130436885 | -0.183960387 | 0.490226326 | 0.396201891 | 0.316382072 |
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0.710883277 | 0.50472972 | 0.37701794 | 0.276436812 | 0.265362095 | 0.214687204 | 0.174849113 |
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0.265288193 | -0.446853384 | 0.196416397 | -0.110376596 | 0.139472173 | -0.13673416 | 0.085656549 |
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0.09057594 | -0.129906618 | -0.173199899 | 0.177854897 | 0.108703018 | -0.085524495 | -0.103253386 |
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-0.070645969 | -0.213980283 | -0.087846741 | -0.389903504 | -0.321747003 | -0.17842853 | -0.021701889 |
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0.039378469 | -0.022086396 | 0.070374143 | 0.022285485 | 0.134276418 | -0.209651757 | 0.058151643 |
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0.139854316 | -0.180680891 | -0.127550531 | 0.157737189 | -0.027411952 | 0.137530567 | -0.134451774 |
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-0.203202698 | 0.278604397 | -0.198153958 | 0.402630535 | -0.109054652 | -0.264792874 | 0.164044114 |
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-0.319474696 | 0.251272882 | -0.008164379 | 0.059050615 | -0.034208937 | 0.225488518 | 0.158104491 |
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0.295981088 | 0.09576584 | 0.048312128 | 0.019723407 | 0.172978909 | 0.06220716 | -0.41905017 |
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0.426919556 | 0.259878689 | -0.007215665 | -0.131144367 | 0.25121175 | -0.129629771 | 0.403591724 |
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0.038959782 | -0.113710926 | -0.227904855 | 0.339050813 | -0.133303008 | -0.064748891 | -0.201696035 |
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-0.469863359 | -0.267676841 | -0.188020288 | -0.317994993 | 0.412756086 | 0.253021631 | 0.111955286 |
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-0.122814373 | 0.056652546 | -0.125361847 | 0.03792394 | 0.095742055 | -0.182302581 | 0.359575483 |
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-0.057154891 | 0.014271255 | 0.303625758 | -0.210415667 | -0.567849769 | -0.191700829 | -0.00332367 |
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-0.266800377 | 0.279797707 | 0.228807449 | -0.14978157 | 0.145371473 | -0.238209022 | -0.413156342 |
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0.069118103 | 0.008227511 | -0.004079106 | -0.18336757 | -0.303043068 | 0.52714368 | 0.090386865 |
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0.178898959 | 0.359957261 | -0.462995625 | -0.293841774 | 0.004054536 | 0.173258047 | -0.103413131 |
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-0.352501894 | -0.193922056 | 0.002786908 | 0.157568267 | 0.04772871 | -0.046397338 | -0.113553524 |
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-0.000619521 | -0.057816313 | 0.271775541 | 0.090085724 | 0.002030737 | -0.003433238 | 0.40989474 |
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-0.02790961 | 0.257813239 | 0.336314946 | 0.217437021 | -0.064780745 | 0.352986111 | 0.03220845 |
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-0.010841335 | 0.086524979 | 0.460243812 | -0.002655612 | 0.309267431 | 0.129809552 | -0.119165889 |
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0.091314647 | -0.276112686 | 0.005762572 | 0.333431445 | -0.07395688 | 0.301989669 | 0.038025431 |
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0.14152623 | 0.133573785 | 0.115042904 | 0.080041817 | 0.055784316 | 0.029292215 | 0.022344327 |
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-0.083499418 | 0.052323475 | -0.106015238 | 0.003344968 | -0.031996278 | 0.03916795 | -0.019457555 |
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-0.178247777 | -0.058497317 | 0.597846907 | -0.255954135 | 0.379366422 | -0.199217481 | 0.221418054 |
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0.08265144 | -0.13083192 | -0.024132742 | 0.014837636 | 0.131660321 | 0.479388229 | 0.2891919 |
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0.244724903 | -0.059011694 | 0.113956101 | -0.074931386 | 0.04577337 | -0.069612504 | -0.732492773 |
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0.050630806 | 0.316234183 | 0.16660119 | 0.299515683 | -0.49781424 | 0.046244883 | 0.219619924 |
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0.296740909 | 0.376559691 | -0.260809485 | -0.089676204 | -0.027290387 | -0.071210438 | 0.140852117 |
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-0.231235089 | -0.168155577 | -0.115371244 | 0.284151752 | 0.361474786 | 0.202917841 | -0.049017145 |
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0.022603597 | 0.420837275 | -0.236893263 | 0.02900181 | 0.436157574 | 0.238920063 | -0.045962367 |
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0.248024801 | -0.355459446 | -0.140327076 | 0.068202164 | 0.014098586 | -0.168841736 | 0.325882917 |
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-0.393037351 | -0.425564191 | -0.418495874 | -0.066368342 | -0.173649977 | -0.04666024 | -0.080329 |
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-0.081626216 | 0.152259274 | -0.192312604 | -0.041856001 | 0.036286384 | -0.306664296 | 0.054524547 |
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-0.115305892 | 0.069986307 | 0.310505175 | 0.23842857 | -0.096262685 | 0.459593261 | -0.180632462 |
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-0.116757716 | 0.126720851 | 0.055407627 | 0.30600813 | 0.081267972 | -0.470097472 | -0.064137815 |
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-0.021197578 | -0.178071358 | 0.100589652 | -0.310374261 | -0.331807871 | 0.142300685 | 0.047913831 |
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0.334353255 | -0.016725971 | -0.026314358 | -0.397995653 | -0.105887039 | 0.100382932 | -0.137085205 |
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-0.184269598 | 0.036899566 | 0.194841372 | 0.163667167 | -0.204455175 | -0.121267207 | -0.156153276 |
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0.428551026 | -0.165166993 | 0.155004291 | 0.142458525 | 0.008969802 | -0.065554124 | 0.111490515 |
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-0.391157106 | 0.255079755 | -0.004834981 | -0.342258191 | -0.175496507 | 0.081476947 | 0.028734251 |
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-0.059743664 | -0.04851249 | 0.20733712 | -0.095515858 | 0.057436279 | -0.015186946 | 0.173384569 |
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0.002458015 | -0.062420243 | -0.039680685 | 0.361422477 | -0.145522506 | 0.02608135 | 0.040083497 |
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0.117972701 | -0.206198025 | 0.0622901 | 0.17118706 | 0.012651195 | 0.069479213 | -0.122746338 |
The
The ANOVAs and subsequent TukeyHDS tests demonstrated that
Summary statistics of morphometric and meristic characters among the
Species |
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FAr | TBLr | AGr | SPLr.l | IFLr.l |
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V | T4r.l |
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Mean | 1.62 | 0.93 | 1.07 | 0.89 | 0.73 | 0.04 | 0.47 | 0.50 | 0.63 | 0.51 | 0.14 | 0.11 | 0.75 | 0.83 | 1.23 | 7.5 | 6.33 | 6 | 7 | 20.33 | 14.17 |
SD | 0.060 | 0.007 | 0.003 | 0.001 | 0.007 | 0.005 | 0.002 | 0.007 | 0.002 | 0.005 | 0.018 | 0.029 | 0.034 | 0.013 | 0.025 | 0.5 | 0.577 | 0 | 0 | 1.155 | 0.289 |
Lower | 1.56 | 0.92 | 1.06 | 0.89 | 0.72 | 0.04 | 0.47 | 0.49 | 0.63 | 0.51 | 0.12 | 0.08 | 0.73 | 0.82 | 1.20 | 7 | 6 | 6 | 7 | 19 | 14 |
Upper | 1.68 | 0.93 | 1.07 | 0.89 | 0.73 | 0.05 | 0.47 | 0.50 | 0.64 | 0.52 | 0.15 | 0.13 | 0.79 | 0.84 | 1.25 | 8 | 7 | 6 | 7 | 21 | 14.5 |
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Mean | 1.67 | 0.98 | 1.11 | 0.94 | 0.71 | 0.15 | 0.52 | 0.57 | 0.69 | 0.57 | 0.18 | 0.16 | 0.79 | 0.87 | 1.29 | 9 | 7.83 | 7.5 | 8.33 | 23.33 | 15 |
SD | 0.099 | 0.014 | 0.017 | 0.005 | 0.013 | 0.022 | 0.016 | 0.042 | 0.028 | 0.012 | 0.013 | 0.047 | 0.017 | 0.007 | 0.003 | 0.5 | 0.289 | 0.5 | 0.577 | 0.577 | 0 |
Lower | 1.55 | 0.96 | 1.09 | 0.93 | 0.70 | 0.12 | 0.51 | 0.54 | 0.67 | 0.56 | 0.17 | 0.12 | 0.78 | 0.86 | 1.29 | 8.5 | 7.5 | 7 | 8 | 23 | 15 |
Upper | 1.74 | 0.99 | 1.13 | 0.94 | 0.72 | 0.17 | 0.54 | 0.62 | 0.72 | 0.58 | 0.20 | 0.21 | 0.81 | 0.87 | 1.29 | 9.5 | 8 | 8 | 9 | 24 | 15 |
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Mean | 1.65 | 0.97 | 0.86 | 0.82 | 0.67 | 0.12 | 0.45 | 0.55 | 0.68 | 0.56 | 0.17 | 0.57 | 0.80 | 0.86 | 1.33 | 7.75 | 6.42 | 5.83 | 7.67 | 21.33 | 14.33 |
SD | 0.025 | 0.012 | 0.008 | 0.037 | 0.022 | 0.044 | 0.022 | 0.021 | 0.025 | 0.022 | 0.023 | 0.025 | 0.020 | 0.017 | 0.060 | 0.418 | 0.376 | 0.258 | 1.211 | 1.366 | 1.033 |
Lower | 1.61 | 0.95 | 0.85 | 0.79 | 0.65 | 0.08 | 0.42 | 0.52 | 0.64 | 0.53 | 0.14 | 0.53 | 0.78 | 0.85 | 1.28 | 7 | 6 | 5.5 | 7 | 20 | 13 |
Upper | 1.68 | 0.98 | 0.87 | 0.89 | 0.71 | 0.18 | 0.49 | 0.57 | 0.70 | 0.59 | 0.21 | 0.60 | 0.83 | 0.89 | 1.44 | 8 | 7 | 6 | 10 | 23 | 15 |
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Mean | 1.69 | 1.03 | 1.15 | 0.99 | 0.78 | 0.21 | 0.47 | 0.54 | 0.68 | 0.66 | 0.19 | 0.33 | 0.78 | 0.81 | 1.35 | 8.17 | 6.5 | 6 | 10 | 22.33 | 13.5 |
SD | 0.096 | 0.001 | 0.016 | 0.006 | 0.005 | 0.031 | 0.030 | 0.003 | 0.011 | 0.005 | 0.011 | 0.006 | 0.004 | 0.001 | 0.032 | 0.289 | 0.5 | 0 | 0 | 0.577 | 1.323 |
Lower | 1.58 | 1.03 | 1.13 | 0.97 | 0.77 | 0.18 | 0.44 | 0.54 | 0.67 | 0.65 | 0.18 | 0.32 | 0.78 | 0.81 | 1.31 | 8 | 6 | 6 | 10 | 22 | 12 |
Upper | 1.75 | 1.03 | 1.16 | 0.99 | 0.78 | 0.25 | 0.50 | 0.54 | 0.69 | 0.66 | 0.20 | 0.34 | 0.78 | 0.81 | 1.38 | 8.5 | 7 | 6 | 10 | 23 | 14.5 |
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Mean | 1.62 | 0.93 | 1.09 | 0.90 | 0.70 | 0.13 | 0.43 | 0.54 | 0.68 | 0.58 | 0.25 | 0.54 | 0.80 | 0.84 | 1.27 | 7.94 | 6.56 | 5.94 | 9.63 | 20.63 | 14.38 |
SD | 0.077 | 0.0357 | 0.009 | 0.014 | 0.029 | 0.048 | 0.034 | 0.044 | 0.021 | 0.031 | 0.046 | 0.016 | 0.028 | 0.017 | 0.021 | 0.320 | 0.496 | 0.177 | 0.518 | 1.061 | 0.835 |
Lower | 1.61 | 0.95 | 0.85 | 0.79 | 0.65 | 0.08 | 0.42 | 0.52 | 0.64 | 0.53 | 0.14 | 0.53 | 0.78 | 0.85 | 1.23 | 7 | 6 | 5.5 | 7 | 20 | 13 |
Upper | 1.68 | 0.987 | 0.87 | 0.89 | 0.71 | 0.18 | 0.49 | 0.57 | 0.70 | 0.59 | 0.21 | 0.60 | 0.83 | 0.89 | 1.44 | 8 | 7 | 6 | 10 | 23 | 15 |
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Mean | 1.62 | 0.97 | 1.07 | 0.91 | 0.70 | 0.17 | 0.46 | 0.48 | 0.65 | 0.50 | 0.18 | 0.23 | 0.73 | 0.83 | 1.26 | 7.75 | 5.67 | 6 | 8.17 | 23.33 | 14.67 |
SD | 0.045 | 0.033 | 0.013 | 0.009 | 0.026 | 0.034 | 0.043 | 0.041 | 0.024 | 0.023 | 0.073 | 0.150 | 0.050 | 0.016 | 0.027 | 0.418 | 0.408 | 0 | 0.983 | 1.633 | 1.033 |
Lower | 1.55 | 0.93 | 1.05 | 0.90 | 0.66 | 0.10 | 0.40 | 0.42 | 0.62 | 0.47 | 0.098 | 0.13 | 0.68 | 0.80 | 1.24 | 7 | 5 | 6 | 7 | 21 | 13 |
Upper | 1.67 | 1.01 | 1.08 | 0.92 | 0.74 | 0.20 | 0.51 | 0.55 | 0.68 | 0.54 | 0.29 | 0.53 | 0.82 | 0.84 | 1.31 | 8 | 6 | 6 | 10 | 26 | 16 |
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Mean | 1.55 | 0.85 | 1.03 | 0.81 | 0.54 | 0.05 | 0.33 | 0.45 | 0.55 | 0.49 | 0.11 | 0.21 | 0.66 | 0.75 | 1.18 | 8 | 6.25 | 6 | 8.25 | 23 | 14 |
SD | 0.083 | 0.033 | 0.009 | 0.037 | 0.049 | 0.068 | 0.017 | 0.033 | 0.087 | 0.033 | 0.007 | 0.015 | 0.032 | 0.032 | 0.020 | 0.408 | 0.645 | 0 | 0.5 | 0.816 | 0.408 |
Lower | 1.48 | 0.81 | 1.02 | 0.76 | 0.49 | -0.03 | 0.31 | 0.41 | 0.44 | 0.44 | 0.11 | 0.19 | 0.62 | 0.72 | 1.16 | 7.5 | 5.5 | 6 | 8 | 22 | 13.5 |
Upper | 1.66 | 0.88 | 1.04 | 0.84 | 0.61 | 0.13 | 0.35 | 0.48 | 0.62 | 0.52 | 0.12 | 0.22 | 0.70 | 0.80 | 1.20 | 8.5 | 7 | 6 | 9 | 24 | 14.5 |
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Mean | 1.56 | 0.83 | 0.83 | 0.82 | 0.65 | -0.03 | 0.41 | 0.47 | 0.61 | 0.48 | 0.10 | 0.29 | 0.67 | 0.78 | 1.20 | 6.75 | 6.29 | 5.63 | 8.67 | 19.25 | 13.25 |
SD | 0.088 | 0.055 | 0.016 | 0.028 | 0.049 | 0.054 | 0.030 | 0.031 | 0.035 | 0.055 | 0.070 | 0.165 | 0.063 | 0.035 | 0.051 | 1.177 | 0.450 | 0.377 | 0.985 | 1.545 | 0.399 |
Lower | 1.41 | 0.72 | 0.80 | 0.77 | 0.58 | -0.13 | 0.35 | 0.42 | 0.55 | 0.36 | 0.02 | 0.12 | 0.55 | 0.73 | 1.12 | 5 | 6 | 5 | 7 | 15 | 12.5 |
Upper | 1.66 | 0.92 | 0.85 | 0.86 | 0.72 | 0.05 | 0.46 | 0.51 | 0.68 | 0.55 | 0.28 | 0.53 | 0.76 | 0.85 | 1.27 | 8 | 7 | 6 | 10 | 21 | 14 |
Significant
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0.007 | |||||||||||
0.00 | 0.005 | 0.022 | |||||||||
0.040 | < 0.001 | 0.001 | 0.001 | 0.023 | |||||||
0.016 | |||||||||||
sp. vs. |
0.021 | 0.001 | < 0.001 | < 0.001 | 0.006 | ||||||
0.005 | 0.00 | < 0.001 | 0.030 | 0.040 | 0.036 | ||||||
0.00 | < 0.001 | 0.017 | |||||||||
0.003 | |||||||||||
< 0.001 | |||||||||||
< 0.001 | 0.002 | ||||||||||
sp. vs. |
0.002 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.005 | < 0.001 | 0.018 | |
< 0.001 | 0.00 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.017 | 0.001 | < 0.001 | 0.023 | ||
0.00 | < 0.001 | 0.002 | |||||||||
0.00 | < 0.001 | ||||||||||
0.00 | < 0.001 | 0.006 | 0.035 | ||||||||
sp. vs. |
< 0.001 | 0.00 | < 0.001 | < 0.001 | 0.001 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | ||
< 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.007 | < 0.001 | < 0.001 | < 0.001 | ||||
0.016 | < 0.001 | 0.001 | 0.030 | ||||||||
<0.001 | 0.006 | ||||||||||
sp. vs. |
< 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.001 | < 0.001 | 0.016 | 0.001 | 0.013 | < 0.001 | |
<0.001 | 0.00 | <0.001 | <0.001 | <0.001 | 0.021 | 0.019 | 0.038 | 0.004 | <0.001 | ||
0.031 | 0.010 | 0.018 | |||||||||
sp. vs. |
0.016 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.002 | < 0.001 | < 0.001 | < 0.001 | 0.012 | |
< 0.001 | 0.00 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.007 | |||
sp. vs. |
< 0.001 | 0.017 | < 0.001 | < 0.001 | 0.013 | < 0.001 | 0.017 | 0.003 | 0.032 | ||
< 0.001 | 0.00 | < 0.001 | < 0.001 | 0.003 | 0.013 | 0.038 | |||||
0.00 | < 0.001 | < 0.001 | |||||||||
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0.008 | < 0.001 | ||||||||||
< 0.001 | |||||||||||
< 0.001 | |||||||||||
0.011 | |||||||||||
sp. vs. |
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0.041 | |||||||||||
0.003 | < 0.001 | ||||||||||
0.026 | < 0.001 | ||||||||||
0.007 | < 0.001 | ||||||||||
< 0.001 | < 0.001 | ||||||||||
sp. vs. |
0.002 | < 0.001 | |||||||||
< 0.001 | < 0.001 | < 0.001 | < 0.001 | 0.015 | |||||||
0.004 | |||||||||||
0.001 | |||||||||||
0.045 | |||||||||||
sp. vs. |
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0.038 | |||||||||||
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0.010 | |||||||||||
sp. vs. |
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0.011 | |||||||||||
0.019 | 0.002 | ||||||||||
sp. vs. |
0.065 | ||||||||||
0.022 | 0.035 | ||||||||||
sp. vs. |
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< 0.001 | |||||||||||
< 0.001 |
Boxplot comparisons of meristic characters among the
Violin plots of the normalized morphometric characters overlain with box plots showing the range, frequency, mean (white dot), and 50% quartile (black rectangle) of characters where interspecific statistical differences were recovered (see Table
Adult male,
Body relatively short (
Forelimbs moderate in stature, relatively short (
Tail 108.4 mm in length, 4.5 mm in width at base, tapering to a point; dorsal scales of flat, square with conical, keeled tubercles in anterior part; median row of transversely expanded subcaudal scales, significantly larger than dorsal caudal scales on original portion; base of tail bearing hemipenal swellings; and postcloacal scales flat, imbricate.
Dorsal views of
The specimens were found at night, between 19:45 and 21:00 h, on the ground in an area along the National Highway 25. The surrounding habitat was secondary montane forest with woody trees. The temperature and humidity were approximately 32.6 °C and 57% (Fig.
Habitat of
The new species is named after the type locality of Gia Lai Province, Central Highlands, Vietnam.
Adult male,
Body relatively short (
Forelimbs moderate in stature, relatively short (
Tail 37.8 mm in length, first 17.1 mm original, 6.1 mm in width at base, tapering to a point; dorsal scales of flat, square with conical, keeled tubercles, regenerated portion covered with small, smooth subcircular scales; median row of transversely expanded subcaudal scales, significantly larger than dorsal caudal scales on original portion; base of tail bearing hemipenal swellings; and postcloacal scales flat, imbricate.
View of
The specific epithet of the new species refers to the type locality of the new species in Muangfuang District, Vientiane Province, Central Laos.
The type series was collected between 19:10 and 19:30 h, on the ground inside Sinxay Temple, at an elevation of 276 m a.s.l. The surrounding habitat was disturbed lowland karst forest (Fig.
Habitat of
Morphological comparisons indicated that
The
For supporting field work and issuing relevant permits, we are grateful to Chu Se Protection Forest’s ranger station and The People’s Committee of Chu Se District, Gia Lai Vietnam, and National University of Laos. We thank Oanh Van Lo and Khanh Quoc Nguyen for his assistance in the field in Vietnam. Many thanks to Thomas Ziegler (Cologne) and Truong Quang Nguyen (Hanoi) for providing comments on the manuscript. Field surveys in Laos were financially supported by Rufford Foundation (ID: 31189-1) to THN and SS. Field surveys in Vietnam were partially funded by the Mohamed bin Zayed Species Conservation Fund (Project Number: 192521666) and the Vietnam National Foundation of Science and Technology Development (NAFOSTED, Grant No. 106.06-2021.28) to VQL. Equipment was supported by IDEA WILD. Research of THN was funded by the Master, PhD Scholarship Programme of Vingroup Innovation Foundation (VINIF), code VINIF.2022.TS125. Research of VQL in the Herpetology Laboratory, Department of Biology, La Sierra University, U.S. was supported by the Fulbright Program.
Measurements and morphological characters of the type series of
tables (Excel spreadsheet)
table S1: Measurements (in mm) and morphological characters of the type series of