Corresponding author: Anchalee Aowphol (
Academic editor: A. Crottini
The taxonomy and geographic distributions of species of crab-eating frogs (
Yodthong S, Stuart BL, Aowphol A (2019) Species delimitation of crab-eating frogs (
Southeast Asia harbors high levels of amphibian species diversity and endemism (
Species of frogs in the genus
Two of these species,
In this study, we examined morphology and mitochondrial DNA variation in historical and newly-collected museum specimens of the
During 2015–2017, specimens of
Map of sampling localities of the
Specimens of
Species identification | Locality | Museum No. | GenBank Acession No. | Type of analyses | Reference | |
---|---|---|---|---|---|---|
Previous study | This study | |||||
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Manila, Luzon, Philippines | – | B | This study | ||
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Malaysia | – | B | This study | |
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Northern Luzon | – | B | This study | |
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Northern Luzon | – | B | This study | |
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Chonburi, Thailand | – | B | This study | |
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Mueang Surat Thani, Surat Thani, Thailand | – | B | This study | |
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Moo Ko Chumphon National Park, Chumphon, Thailand | – | B | This study | |
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Moo Ko Chumphon National Park, Chumphon, Thailand | – | B | This study | |
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Moo Ko Chumphon National Park, Chumphon, Thailand | – | B | This study | |
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Ko Libong, Trang, Thailand | – | B | This study | |
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Songkhla lake, Songkhla, Thailand | – | B | This study | |
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Songkhla lake, Phatthalung, Thailand | – | B | This study | |
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Kleang, Rayong, Thailand | – | B | This study | |
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Kleang, Rayong, Thailand | – | B | This study | |
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Kleang, Rayong, Thailand | – | B | This study | |
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Kleang, Rayong, Thailand | – | B | This study | |
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Mueang Trat, Trat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Suk Samran, Ranong, Thailand | – | B | This study | |
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Suk Samran, Ranong, Thailand | – | B | This study | |
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Suk Samran, Ranong, Thailand | – | B | This study | |
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Sam Roi Yot, Prachuap Khiri Khan, Thailand |
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A | This study | |
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Sam Roi Yot, Prachuap Khiri Khan, Thailand |
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A, B | This study | |
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Sam Roi Yot, Prachuap Khiri Khan, Thailand |
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A | This study | |
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Sam Roi Yot, Prachuap Khiri Khan, Thailand | – | B | This study | |
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Kraburi, Ranong, Thailand |
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A, B | This study | |
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Kraburi, Ranong, Thailand |
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A, B | This study | |
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Mueang, Phuket, Thailand | – | B | This study | |
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Mueang, Phuket, Thailand |
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A | This study | |
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Mueang, Phuket, Thailand |
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A, B | This study | |
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Ko Samui, Surat Thani, Thailand |
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A, B | This study | |
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Ko Samui, Surat Thani, Thailand | – | B | This study | |
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Ko Samui, Surat Thani, Thailand |
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A, B | This study | |
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Mueang Phang-nga, Phang-nga, Thailand |
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A, B | This study | |
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Mueang Phang-nga, Phang-nga, Thailand |
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A, B | This study | |
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Mueang Phang-nga, Phang-nga, Thailand |
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A, B | This study | |
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Mueang Phang-nga, Phang-nga, Thailand | – | B | This study | |
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Mueang Phuket, Phuket, Thailand |
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A, B | This study | |
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Mueang Phuket, Phuket, Thailand | – | B | This study | |
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Mueang Phuket, Phuket, Thailand | – | B | This study | |
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Ko Lanta, Krabi, Thailand |
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A, B | This study | |
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Ko Lanta, Krabi, Thailand | – | B | This study | |
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Ko Lanta, Krabi, Thailand |
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A | This study | |
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Ko Lanta, Krabi, Thailand |
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A | This study | |
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Khanom, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Ko Chang, Trat, Thailand |
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A, B | This study | |
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Ko Chang, Trat, Thailand |
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A, B | This study | |
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Ko Chang, Trat, Thailand |
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A, B | This study | |
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Ko Chang, Trat, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand |
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A, B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand |
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A, B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand |
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A, B | This study | |
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Kraburi, Ranong, Thailand |
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A, B | This study | |
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Kraburi, Ranong, Thailand | – | B | This study | |
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Mueang Krabi, Krabi, Thailand | – | B | This study | |
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Mueang Krabi, Krabi, Thailand | – | B | This study | |
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Kui Buri, Prachuap Khiri Khan, Thailand | – | B | This study | |
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La-ngu, Satun, Thailand |
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A, B | This study | |
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La-ngu, Satun, Thailand | – | B | This study | |
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La-ngu, Satun, Thailand |
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A, B | This study | |
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La-ngu, Satun, Thailand |
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A, B | This study | |
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Kleang, Rayong, Thailand |
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A, B | This study | |
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Kleang, Rayong, Thailand |
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A, B | This study | |
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Manila, Philippines | – |
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A |
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Negros Island, Philippines | – |
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A |
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Hainan, China | – |
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A |
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Dacope, Khulna, Bangladesh | – |
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A |
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Teknaf, Cox’s Bazar, Bangladesh | – |
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A |
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Cianjur, Java, Indonesia | – |
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A |
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Padang, Sumatra, Indonesia | – |
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A |
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Selangor, Malaysia | – |
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A |
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Bogor, Java, Indonesia | – |
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A |
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Banyumas, Java, Indonesia | – |
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A |
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Malang, East Java, Indonesia | – |
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A |
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Denpasar, Bali, Indonesia | – |
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A |
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Cianjur, Java, Indonesia | – | B | This study | ||
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Java, Indonesia | – | B | This study | |
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Java, Indonesia | – | B | This study | |
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Java, Indonesia | CMNH 161102 | – | B | This study |
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Java, Indonesia | – | B | This study | |
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Java, Indonesia | – | B | This study | |
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Java, Indonesia | – | B | This study | |
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Nakhon Si Thammarat, Thailand | – | B | This study | |
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Phatthalung, Thailand | – | B | This study | |
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Phatthalung, Thailand | – | B | This study | |
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Phatthalung, Thailand | – | B | This study | |
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Phatthalung, Thailand | – | B | This study | |
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Phatthalung, Thailand | – | B | This study | |
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Songkhla, Thailand | – | B | This study | |
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Songkhla, Thailand | – | B | This study | |
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Nong Chick, Pattani, Thailand | – | B | This study | |
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Su-Ngai Kolok, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
|
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
|
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Tak Bai, Narathiwat, Thailand | – | B | This study | |
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Pak Phayun, Phatthalung, Thailand | – | B | This study | |
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Pak Phayun, Phatthalung, Thailand | – | B | This study | |
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Pak Phayun, Phatthalung, Thailand | – | B | This study | |
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Pak Phayun, Phatthalung, Thailand | – | B | This study | |
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Pak Phayun, Phatthalung, Thailand | – | B | This study | |
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Su-Ngai Kolok, Narathiwat, Thailand | – | B | This study | |
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Nong Chick, Pattani, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Khuan Khanun, Phatthalung, Thailand |
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A | This study | |
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Khuan Khanun, Phatthalung, Thailand |
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A, B | This study | |
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Khuan Khanun, Phatthalung, Thailand | – | B | This study | |
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Khuan Khanun, Phatthalung, Thailand |
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A | This study | |
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Khuan Khanun, Phatthalung, Thailand |
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A, B | This study | |
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Khuan Khanun, Phatthalung, Thailand | – | B | This study | |
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Khuan Khanun, Phatthalung, Thailand |
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A, B | This study | |
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Khuan Khanun, Phatthalung, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand |
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A, B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand |
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A, B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
|
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Pak Phanang, Nakhon Si Thammarat, Thailand |
|
A, B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
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Pak Phanang, Nakhon Si Thammarat, Thailand | – | B | This study | |
Pelabuhan ratu, Java, Indonesia | – |
|
A |
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Makassar, Sulawesi, Indonesia | – |
|
A |
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Makassar, Sulawesi, Indonesia | – |
|
A |
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Selatan, Sulawesi, Indonesia | – |
|
A |
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Malang, Java, Indonesia | – |
|
A |
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Java, Indonesia | – |
|
A |
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Yunan, China | – |
|
A |
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Quezon, Luzon Island, Philippines | – |
|
A |
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Mangalore, India | – |
|
A |
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Surat Thani, Thailand | – |
|
A |
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Kuala Lumpur, Malaysia | – |
|
A |
|
Total genomic DNA was extracted from liver or muscle tissue using the GF-1 Tissue DNA Extraction Kit (Vivantis Inc.). A 961–962 bp fragment of mitochondrial (mt) DNA that encodes part of the 16S rRNA gene was amplified by the polymerase chain reaction (PCR; 94 °C 45s, 58 °C 30s, 72 °C 1 min) for 35 cycles using the primer pairs L16SRanaIII (
Homologous sequences of
Morphological analyses were performed on 108 sexually mature individuals (61 males, 47 females) of
Measurements were taken with digital Vernier calipers to the nearest 0.1 mm. Twenty-three morphological characters were measured following
Qualitative characters were taken on the presence and condition of the vomerine ridge, skin on dorsum, coloration and pattern on dorsum, vocal sac pigmentation, fejervaryan lines (conspicuous ventrolateral lines on the ventral side of the body), tubercles on forelimbs and hindlimbs, dermal fringe on fingers II and III, inner tarsal ridge, dermal flap on outer side of Toe V, and foot webbing.
To correct for body size, each mensural character was divided by
The aligned dataset contained 61 individuals and 981 characters. The standard deviation of split frequencies was 0.003331 among the two Bayesian runs, and the Estimated Sample Sizes (ESS) of parameters were ≥ 200. The Bayesian analysis recovered the
Uncorrected pairwise sequence divergences (
Bayesian consensus phylogram of the mitochondrial16S rRNA gene of
Principal component analysis of morphological measurements from males (
Uncorrected pairwise sequence divergences (
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– | |||||||
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(12.8) | – | ||||||
12.8 | ||||||||
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(12.1–12.7) | (0.2–0.4) | (0.9) | |||||
12.4 | 0.3 | 0.9 | ||||||
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(16.2) | (12.2) | (11.7–13.5) | – | ||||
16.2 | 12.2 | 12.6 | ||||||
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(17.8–18.2) | (13.7–13.9) | (13.4–15.0) | (11.4–12.3) | (0.0–1.6) | |||
18.0 | 13.8 | 13.9 | 11.5 | 0.4 | ||||
|
(14.3–18.6) | (13.9–14.2) | (13.4–14.8) | (9.5–12.9) | (0.9–3.4) | (0.0–1.6) | ||
17.4 | 14.1 | 14.0 | 11.7 | 1.7 | 0.4 | |||
|
(12.5–17.1) | (13.9–14.3) | (13.4–15.1) | (10.7–12.8) | (8.8–10.7) | (8.3–11.1) | (0.0–1.4) | |
15.5 | 13.7 | 14.1 | 11.9 | 9.7 | 9.3 | 0.3 | ||
|
(10.9–12.5) | (12.8–13.7) | (12.3–13.5) | (9.5–10.2) | (9.8–11.0) | (8.9–11.0) | (4.5–7.9) | (0.6–6.0) |
11.7 | 13.4 | 13.2 | 9.8 | 10.5 | 9.3 | 6.5 | 3.6 |
Factor loading on the first three principal components of 23 morphological characters for male and female
Character | Males | Females | ||||
---|---|---|---|---|---|---|
|
0.395 | -0.829 | -0.211 | 0.136 | -0.921 | -0.008 |
rHL | 0.614 | 0.462 | -0.306 | 0.628 | 0.230 | 0.254 |
rHW | 0.795 | -0.093 | -0.262 | 0.660 | -0.445 | 0.297 |
rSTL | 0.623 | 0.610 | -0.222 | 0.762 | 0.201 | 0.353 |
rNS | 0.272 | 0.574 | -0.157 | 0.640 | 0.252 | 0.213 |
rSL | 0.725 | 0.300 | -0.343 | 0.829 | -0.119 | 0.162 |
rNTL | 0.511 | 0.703 | -0.155 | 0.654 | 0.165 | 0.310 |
rEN | 0.601 | 0.108 | -0.347 | 0.715 | 0.016 | 0.240 |
rTEL | 0.376 | -0.236 | -0.255 | 0.329 | -0.589 | -0.243 |
rTD | -0.211 | 0.744 | -0.213 | 0.199 | 0.570 | 0.149 |
rIN | 0.166 | -0.041 | -0.319 | 0.562 | -0.185 | 0.136 |
rEL | -0.279 | 0.767 | 0.100 | 0.064 | 0.820 | 0.128 |
rIOD | -0.278 | 0.659 | 0.431 | 0.055 | 0.628 | -0.356 |
rUEW | 0.132 | 0.176 | -0.556 | 0.104 | 0.286 | 0.575 |
rHAL | 0.549 | 0.358 | 0.487 | 0.701 | 0.285 | -0.447 |
rFAL | 0.157 | 0.408 | 0.538 | 0.422 | 0.074 | -0.325 |
rTHIGHL | 0.768 | -0.128 | -0.138 | 0.675 | -0.136 | 0.146 |
rTL | 0.815 | -0.384 | -0.106 | 0.760 | -0.281 | 0.208 |
rFOL | 0.775 | -0.011 | 0.421 | 0.800 | 0.055 | -0.250 |
rTFOL | 0.766 | -0.249 | 0.382 | 0.766 | -0.164 | -0.135 |
r1FL | 0.481 | -0.163 | 0.664 | 0.657 | 0.110 | -0.578 |
rIMTL | 0.436 | 0.034 | 0.252 | 0.478 | 0.089 | -0.383 |
rITL | 0.674 | -0.029 | 0.466 | 0.758 | 0.064 | -0.373 |
Elegenvalue | 6.807 | 4.420 | 2.860 | 8.112 | 3.337 | 2.146 |
Percentage of variance | 29.595 | 19.218 | 12.435 | 35.268 | 14.508 | 9.331 |
Cumulative proportion | 29.595 | 48.813 | 61.248 | 35.268 | 49.776 | 59.107 |
Summary statistics of morphological characters of adult males and females are shown in Table
Comparisons of body sizes of
Characters | Males | Females | ||||||
---|---|---|---|---|---|---|---|---|
|
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|
|
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|
71.3 ± 5.6 | 51.4 ± 5.4 | -13.826 | < 0.0001* | 94.2 ± 6.5 | 69.0 ± 10.1 | 0a | < 0.0001* |
(60.2–79.8) | (42.7–62.7) | (85.1–107.1) | (50.0–81.8) | |||||
rHL | 40.8 ± 1.8 | 39.9 ± 1.7 | 340a | 0.0692 | 39.6 ± 2.4 | 39.4 ± 1.7 | 201.5a | 0.4953 |
(36.7– 43.5) | (37.2– 44.5) | (35.2–42.9) | (35.9–42.2) | |||||
rHW | 37.1 ± 1.8 | 34.6 ± 1.1 | -6.553 | < 0.0001* | 38.2 ± 1.6 | 35.6 ± 1.9 | 65.5a | 0.0001* |
(32.5–40.6) | (32.4–37.1) | (35.0–41.0) | (32.5–38.7) | |||||
rSTL | 30.3 ± 1.2 | 30.1 ± 1.0 | 414.500a | 0.4609 | 29.9 ± 1.1 | 29.3 ± 1.0 | -1.867 | 0.0684 |
(27.5–32.1) | (28.5–32.0) | (27.9–31.6) | (27.6–31.3) | |||||
rNS | 7.2 ± 0.6 | 7.4 ± 0.5 | 1.473 | 0.1460 | 7.3 ± 0.8 | 7.1 ± 0.6 | 177a | 0.2115 |
(5.9–8.5) | (6.1– 8.8) | (5.4–8.2) | (6.1–8.5) | |||||
rSL | 17.0 ± 1.0 | 16.4 ± 0.7 | 262.500a | 0.0033* | 17.2 ± 0.9 | 16.1 ± 0.8 | 88a | 0.0008* |
(14.7–18.2) | (15.2–17.9) | (15.2–18.3) | (14.6–17.9) | |||||
rNTL | 23.28 ± 0.9 | 23.4 ± 1.1 | 0.402 | 0.6893 | 23.0 ± 0.7 | 22.7 ± 0.9 | -0.783 | 0.4375 |
(21.6–25.3) | (21.5–25.8) | (21.8 – 24.0) | (21.2–24.4) | |||||
rEN | 9.5 ± 0.5 | 8.8 ± 0.8 | -3.759 | 0.0004* | 9.5 ± 0.4 | 8.8 ± 0.7 | 95a | 0.0015* |
(8.4–10.7) | (7.4 – 11.3) | (8.5–10.1) | (7.5–10.1) | |||||
rTEL | 3.83 ± 0.64 | 3.23 ± 0.62 | -3.840 | 0.0003* | 4.83 ± 0.64 | 4.28 ± 0.83 | -2.295 | 0.0265* |
(2.73 – 5.21) | (2.40 – 5.05) | (4.06 – 6.32) | (2.76 – 5.98) | |||||
rTD | 7.2 ± 0.5 | 7.9 ± 0.6 | 4.840 | < 0.0001* | 6.9 ± 0.4 | 7.1 ± 0.6 | 1.020 | 0.3131 |
(6.4–8.0) | (6.8– 9.3) | (6.2–7.9) | (5.7– 8.0) | |||||
rIN | 4.9 ± 0.4 | 4.9 ± 0.6 | 371a | 0.1750 | 4.9 ± 0.4 | 4.6 ± 0.4 | -2.270 | 0.0281* |
(4.17 – 5.62) | (3.8–6.2) | (4.3–6.0) | (3.8–5.5) | |||||
rEL | 9.8 ± 0.7 | 11.5 ± 1.1 | 7.026 | < 0.0001* | 8.9 ± 0.9 | 10.1 ± 1.0 | 3.850 | 0.0004* |
(8.1–11.2) | (9.2–13.4) | (7.5–10.3) | (8.3–12.4) | |||||
rIOD | 4.8 ± 0.6 | 6.2 ± 0.7 | 7.902 | < 0.0001* | 5.0 ± 0.4 | 5.6 ± 0.7 | 3.158 | 0.0028* |
(3.7 – 5.9) | (4.6– 8.2) | (4.1–5.51) | (4.1–7.5) | |||||
rUEW | 8.3 ± 0.7 | 8.2 ± 0.6 | -0.270 | 0.7882 | 8.0 ± 0.8 | 7.9 ± 0.7 | -0.140 | 0.8895 |
(6.8–9.6) | (7.1–9.5) | (6.4–9.4) | (6.3–9.2) | |||||
rHAL | 24.6 ± 0.9 | 24.7 ± 1.1 | 0.081 | 0.9359 | 23.9 ± 1.2 | 24.0 ± 1.5 | 218a | 0.7692 |
(23.2–26.3) | (21.3–26.8) | (21.2–25.6) | (21.3–27.45) | |||||
rFAL | 19.3 ± 0.9 | 19.8 ± 1.2 | 1.609 | 0.1130 | 18.7 ± 0.8 | 18.8 ± 1.3 | 0.084 | 0.9335 |
(17.6–21.4) | (17.8– 22.5) | (17.4–20.0) | (16.7–21.4) | |||||
rTHIGHL | 47.8 ± 1.93 | 45.5 ± 1.9 | 178.5a | < 0.0001* | 46.0 ± 2.4 | 43.6 ± 2.1 | 92a | 0.0012 |
(42.1–51.1) | (42.6–49.3) | (40.0–48.5) | (39.9–47.6) | |||||
rTL | 52.0 ± 1.4 | 47.6 ± 2.2 | 53a | < 0.0001* | 50.8 ± 3.0 | 46.4 ± 2.3 | 62.5a | < 0.0001* |
(48.7–55.6) | (41.0–53.0) | (42.7–54.13) | (43.4 – 50.7) | |||||
rFOL | 54.1 ± 2.2 | 51.8 ± 3.0 | 261.5a | 0.0027* | 52.1 ± 1.5 | 50.8 ± 3.1 | 174a | 0.1842 |
(49.8–57.8) | (43.4–58.3) | (50.0–55.0) | (44.1–55.2) | |||||
rTFOL | 79.6 ± 3.3 | 75.1 ± 4.0 | 162a | < 0.0001* | 77.6 ± 4.0 | 72.9 ± 4.6 | 109.5a | 0.004616* |
(73.6–86.5) | (63.6– 81.6) | (71.4–87.0) | (66.3–81.2) | |||||
r1FL | 18.9 ± 1.3 | 18.2 ± 1.3 | -1.967 | 0.0539 | 19.0 ± 1.1 | 18.9 ± 1.1 | -0.173 | 0.8637 |
(17.2–21.2) | (16.2–20.8) | (16.6–20.6) | (17.0–21.0) | |||||
rIMTL | 6.0 ± 0.6 | 5.8 ± 0.6 | 342.5a | 0.0773 | 5.9 ± 0.5 | 6.0 ± 0.5 | 252a | 0.6317 |
(4.0–6.9) | (4.2–7.0) | (4.7–6.5) | (4.8–6.7) | |||||
rITL | 18.6 ± 1.1 | 17.9 ± 1.8 | 327.5a | 0.0469* | 18.4 ± 1.1 | 18.1 ± 1.4 | -0.901 | 0.3724 |
(15.1–20.1) | (14.8–21.8) | (15.9–19.8) | (15.2–20.7) | |||||
1.1 ± 0.0 | 1.2 ± 0.0 | 4.913 | < 0.0001* | 1.0 ± 0.0 | 1.1 ± 0.1 | 4.462 | < 0.0001* | |
(1.0–1.2) | (1.1–1.2) | (1.0–1.1) | (1.0–1.2) | |||||
0.1 ± 0.0 | 0.2 ± 0.0 | 10.343 | < 0.0001* | 0.1 ± 0.0 | 0.2 ± 0.0 | 4.619 | < 0.0001* | |
(0.1–0.2) | (0.1–0.2) | (0.1– 0.2) | (0.1– 0.2) | |||||
0.4 ± 0.0 | 0.4 ± 0.0 | -1.448 | 0.1529 | 0.43 ± 0.0 | 0.4 ± 0.0 | -4.426 | < 0.0001* | |
(0.4–0.5) | (0.4–0.5) | (0.4–0.5) | (0.4–0.5) | |||||
0.2 ± 0.0 | 0.3 ± 0.0 | 8.662 | < 0.0001* | 0.2 ± 0.0 | 0.3 ± 0.0 | 375a | 0.0008* | |
(0.2–0.3) | (0.2–0.3) | (0.2–0.3) | (0.2–0.3) | |||||
0.8 ± 0.1 | 0.8 ± 0.1 | 4.505 | < 0.0001* | 0.8 ± 0.1 | 0.8 ± 0.1 | 2.278 | 0.0275* | |
(0.6–0.9) | (0.7–1.0) | (0.6–0.9) | (0.7–0.9) | |||||
0.6 ± 0.0 | 0.7 ± 0.1 | 8.775 | < 0.0001* | 0.5 ± 0.1 | 0.6 ± 0.1 | 5.664 | < 0.0001* | |
(0.5–0.7) | (0.6–0.8) | (0.5–0.6) | (0.5–0.8) | |||||
1.0 ± 0.1 | 1.3 ± 0.2 | 9.196 | < 0.0001* | 0.9 ± 0.1 | 1.1 ± 0.1 | 6.303 | < 0.0001* | |
(0.9–1.2) | (1.0–1.6) | (0.8–1.1) | (1.0–1.4) | |||||
1.0 ± 0.2 | 0.8 ± 0.1 | -6.372 | < 0.0001* | 1.0 ± 0.1 | 0.8 ± 0.1 | -3.839 | 0.0004* | |
(0.8–1.5) | (0.5–1.2) | (0.8–1.1) | (0.6–1.3) | |||||
0.7 ± 0.1 | 0.7 ± 0.1 | -3.201 | 0.0022* | 0.8 ± 0.1 | 0.7 ± 0.1 | -2.754 | 0.0085* | |
(0.6–0.9) | (0.6–0.8) | (0.6–0.9) | (0.6–0.9) | |||||
0.4 ± 0.1 | 0.3 ± 0.1 | -6.585 | < 0.0001* | 0.6 ± 0.1 | 0.4 ± 0.1 | 108a | 0.0044* | |
(0.3–0.6) | (0.2–0.4) | (0.4–0.8) | (0.2–0.6) | |||||
0.8 ± 0.0 | 0.8 ± 0.1 | 1.385 | 0.1712 | 0.8 ± 0.0 | 0.8 ± 0.1 | 0.031 | 0.9754 | |
(0.7–0.9) | (0.7–0.9) | (0.7–0.8) | (0.7– 0.9) | |||||
0.9 ± 0.0 | 1.0 ± 0.0 | 741a | < 0.0001* | 0.9 ± 0.1 | 0.9 ± 0.0 | 2.574 | 0.0134* | |
(0.9–1.0) | (0.9–1.1) | (0.8–1.0) | (0.9–1.0) | |||||
1.0 ± 0.0 | 1.1 ± 0.1 | 755a | < 0.0001* | 1. ± 0.1 | 1.01 ± 0.1 | 405a | < 0.0001* | |
(1.0–1.1) | (0.9–1.2) | (1.0–1.2) | (1.0–1.2) | |||||
0.1 ± 0.0 | 0.1 ± 0.0 | 615.5a | 0.0296 | 0.1 ± 0.0 | 0.1 ± 0.0 | 3.342 | 0.0017* | |
(0.1–0.1) | (0.1–0.2) | (0.1–0.1) | (0.1–0.2) |
The genetic and morphometric data provide congruent, independent lines of evidence to support the hypothesis that
Dubios and Ohler (2000) designated and described the neotype adult male,
Adult male neotype of
Forelimbs short, rather stout [rather thin according to
Hindlimbs moderately short, robust; tibia longer than thigh, but shorter than distance from base of inner metatarsal tubercle to tip of Toe IV; toes long, thin; tips of toes rounded [pointed according to
Plantar and metatarsal views of
Skin on snout and interorbital region shagreen; skin on eyelid with glandular warts and spinules; skin on dorsum with irregular skin folds, with intervening glandular warts and spinules; dorsolateral fold extending posteriorly to two-thirds length of dorsum; skin on side of head with small spinules; skin on flank with glandular warts; skin on cloacal region with dense glandular warts; skin on forelimbs, thigh, tibia and tarsus with glandular warts and spinules; skin on ventral surfaces smooth, except dense, fine spinules on chin. Nuptial pad with small translucent spinules on dorsal and medial surface of Finger I from base of distal phalanx to slightly over the base of prepollax; vocal sac present on both sides of throat, with wrinkled skin covered by triangular dark brown blotches. Fejervaryan lines absent.
Dorsum and side of head medium brown with indistinct dark brown markings; dark brown band between outer margins of upper eyelids; tympanum brown with inferior half more translucent, lighter in coloration than head; flank creamy white with dark brown marbling; three wide dark brown vertical spots on upper lips; wide light brown mid-dorsal stripe continuous from tip of snout to vent; dorsal surfaces of forelimbs, thigh, tibia, and foot brown with dark brown transverse spots; posterior surface of thighs with irregular pattern of dark brown marbling on white background; chin mottled dark brown, throat with triangular dark brown blotches on each side; chest, belly and ventral surfaces of hindlimbs creamy white with indistinct dark brown mottling; ventral surfaces of forelimbs creamy white; ventral surfaces of hand and foot brown; lower lip creamy white with dark brown spots.
Adult male
Adult male
Females are distinctly larger in size (Table
The examined male and female specimens closely resemble the neotype in morphology, with most observed variation pertaining to coloration. Dorsal coloration in preservative varied from medium to very dark brown with darker markings. Markings or spots on dorsum, and transverse spots on dorsal surface of forelimbs and hindlimbs fainter than neotype in some individuals. Flank pale brown with dark brown marbling in some individuals. Ventral coloration pale brown in some individuals, with dark mottling on chin and chest. Ventral surface of hand pale brown or creamy white in some individuals. Dorsal vertebral stripe present (
Based on a combination of the morphological and genetic studies of
Specimens were collected in Thailand (Khuan Khanun District, Phatthalung Province and Pak Panang District, Nakhon Si Thammarat Province) at night (1900–2200 h) following light rain during May and October 2016. At Khuan Khanun, frogs were sampled in grasslands, rice paddy fields near standing or slow flowing ditches, and ponds at 1–24 m elevation (Fig.
Exemplar habitats in Thailand of
Adult female holotype of
Forelimbs short, rather robust; fingers rather long, slightly swollen; tips of fingers slightly rounded, terminus slightly swollen but not expanded into discs; relative finger lengths II < IV < I < III [first finger longer than second and fourth according to
Hindlimbs moderately short, robust; tibia slightly longer than thigh, but shorter than distance from base of inner metatarsal tubercle to tip of Toe IV; toe long, stout; tips of toes rounded, not expanded into discs; relatively toe lengths I < II < III < IV, webbing moderate, deeply excised between toes, formula I1–11/2II1–1III1–2IV2–1V, Toe I webbed to base of distal phalanx; preaxial side of Toe II webbed to point between distal subarticular tubercle and distal phalanx, continuing as narrow fringe to base of distal phalanx; postaxial side of Toe II webbed to base of distal phalanx; preaxial side of Toe III webbed to distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, postaxial side of Toe III webbed to base of distal phalanx; preaxial side of Toe IV wedded to webbed to proximal distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, postaxial side of Toe IV wedded to webbed to proximal distal subarticular tubercle, continuing as narrow fringe to base of distal phalanx, Toe V webbed to base of distal phalanx; dermal flap well developed, extending along postaxial side of Toe V from level of inner metatarsal tubercles to distal phalanx; subarticular tubercles prominent, inner metatarsal tubercle prominent, oval, length about 30% that of Toe I; distinct dermal ridge extending along inner metatarsal tubercle to distal phalanx of Toe I; indistinct inner tarsal ridge on distal two-third of tarsus (Fig.
Skin on snout and between the eyes shagreened; skin on eyelid shagreened with glandular warts; skin on dorsum shagreened with glandular warts and irregular skin folds; dorsolateral fold extending posteriorly to two-thirds length of dorsum; skin on side of head smooth; skin on flank with glandular warts; skin on cloacal region with glandular warts; forelimbs shagreened; thigh with indistinct glandular warts; tibia, tarsus, throat, chest and belly smooth.
Coloration mostly lost in preservative. Dorsum and side of head medium brown with a few dark brown markings; tympanum translucent brown with pale brown spot in center; flank pale brown with faint brown marbling; three wide brown vertical spots on upper lips; dorsal surfaces of forelimbs, thigh, tibia, and foot medium brown with a few dark brown spots, posterior surface of thigh with irregular pattern of indistinct dark brown marbling on light background; chin, chest, belly, and ventral surfaces of forelimb and hindlimb pale brown; ventral surfaces of hand and foot pale brown; lower lip pale brown with a few dark brown spots; vertebral and tibial stripes absent; Fejervaryan lines absent.
Adult male
Adult male
Vomerine ridges slightly closer to choanae than to each other in some individuals. Most adult males have nuptial pads with small translucent spinules on dorsal and medial surface of Finger I from base of distal phalanx to base of prepollax, but some individuals have the nuptial pad extending to slightly over the base of prepollex. Most adult males have dense, fine spinules covering only the chin, but some individuals have dense, fine spinules on the chin and chest. Adult males have vocal sac present on each side of throat with wrinkled skin covered by triangular, very dark brown blotches. Adult males with larger spinules and glandular warts on dorsum, dorsal surfaces of forelimbs, flank, hindlimbs and vent region. Females are distinctly larger in size (Table
Dorsal coloration in preservative varies in males and females from brown to dark brown with darker markings. Two female specimens from Trat Province, Thailand (
Based on a combination of morphological and genetic studies of
In Thailand, specimens were collected at night (1900–2200 h) in a variety of coastal habitats at elevations ranging from 0–16 m asl. Most specimens were observed in marshes near slow flowing ditches, ponds, or canals in mangrove forest (Fig.
Twelve species of
Our study clarifies that two species of crab-eating frogs (
This study provides a basis for revising the identifications of historical and contemporary records (both museum vouchers and literature descriptions) of crab-eating frogs to improve the finer-scale details of the geographic ranges, as well as the natural histories, of
This research was supported by the Center of Excellence on Biodiversity (BDC), Office of Higher Education Commission, Thailand (BDC-PG4-160022). The research protocol was approved by the Institutional Animal Care and Use Committee, Kasetsart University, Thailand (project number OACKU00459). Alan Resetar (
Specimens examined.
Morphological measurements (mm) of adult male specimens of
Characters | ||||
---|---|---|---|---|
|
66.9 | 74.6 ± 3.8 (71.4 – 79.8) | 71.0 ± 5.7 (60.2–78.9) | 51.4 ± 5.4 (42.7–62.7) |
|
25.6 | 29.7 ± 1.1 (28.6 – 31.0) | 29.1 ± 2.1 (24.5–32.4) | 20.5 ± 1.8 (17.3–25.0) |
|
23.7 | 26.2 ± 1.6 (24.37 – 27.9) | 26.6 ± 2.3 (22.0–30.5) | 17.7 ± 1.8 (14.4–22.1) |
|
19.1 | 22.0 ± 0.6 (21.3– 22.8) | 21.56 ± 1.4 (18.4–23.7) | 15.4 ±1.4 (13.3–18.8) |
|
4.3 | 4.9 ± 0.2 (4.7–5.2) | 5.2 ± 0.4 (4.5–6.0) | 3.8 ± 0.40 (3.1–4.7) |
|
10.5 | 11.7 ± 0.7 (11.0–12.6) | 12.2 ± 0.7 (10.8–14.1) | 8.4 ± 0.8 (6.8–10.2) |
|
14.8 | 17.1 ± 0.78 (16.2–18.1) | 16.6 ± 1.2 (14.2– 18.2) | 12.0 ± 1.0 (10.5 –14.4) |
|
6.0 | 6.9 ± 0.5 (6.6–7.6) | 6.7 ± 0.5 (5.6–7.6) | 4.5 ± 0.4 (3.6–5.2) |
|
2.3 | 3.2 ± 0.2 (3.08 – 3.4) | 2.7 ± 0.6 (1.9–3.7) | 1.7 ± 0.4 (1.2–2.8) |
|
5.1 | 5.2 ± 0.5 (4.6–5.7) | 5.1 ± 0.4 (4.2–6.0) | 4.1 ± 0.4 (3.3–4.9) |
|
2.8 | 3.9 ± 0.1 (3.8–3.9) | 3.5 ± 0.4 (2.9–4.4) | 2.5 ± 0.4 (1.9–3.5) |
|
6.6 | 6.9 ± 0.6 (6.1–7.4) | 7.0 ± 0.7 (5.5–8.6) | 5.9 ± 0.6 (4.8–7.1) |
|
3.0 | 3.4 ± 0.2 (3.1–3.6) | 3.5 ± 0.5 (2.7– 4.4) | 3.2 ± 0.4 (2.5–3.8) |
|
4.9 | 6.4 ± 0.4 (6.07 – 6.83) | 5.9 ± 0.7 (4.9–7.3) | 4.2 ± 0.5 (3.2–5.1) |
|
16.2 | 18.00 ± 0.9 (16.9–19.1) | 17.5 ± 1.2 (15.5–19.5) | 12.7 ± 1.4 (10.6–15.7) |
|
13.1 | 15.0 ± 0.9 (14.1–16.0) | 13.6 ± 0.9 (11.5–15.0) | 10.2 ± 1.2 (8.2–12.2) |
|
30.2 | 34.7 ± 2.8 (31.6–38.0) | 34.1 ± 2.8 (29.1–39.1) | 23.4 ± 2.7 (19.0–29.3) |
|
34.5 | 38.4 ± 2.1 (36.5–41.4) | 37.0 ± 3.0 (30.7 – 42.7) | 24.44 ± 2.73 (19.84 – 30.3) |
|
37.2 | 39.8 ± 1.0 (38.6–40.7) | 38.35 ± 2.5 (30.7–43.3) | 26.6 ± 3.1 (21.5–32.9) |
|
56.3 | 59.8 ± 1.4 (58.4–61.3) | 56.2 ± 4.1 (48.7–62.8) | 38.6 ± 4.6 (30.4–48.1) |
|
13.8 | 14.3 ± 0.8 (13.3–15.0) | 13.3 ± 1.1 (11.6–16.1) | 9.4 ± 1.3 (7.2–12.4) |
|
4.1 | 3.9 ± 0.5 (3.5– 4.5) | 4.3 ± 0.4 (3.2–5.2) | 3.0 ± 0.5 (1.9–4.0) |
|
12.4 | 13.2 ± 1.3 (11.3–14.3) | 13.3 ± 1.0 (11.9–14.9) | 9.2 ± 1.4 (7.0–11.3) |
1.1 | 1.1 ± 0.0 (1.1–1.2) | 1.1 ± 0.0 (1.0–1.2) | 1.2 ± 0.0 (1.1–1.2) | |
0.1 | 0.1 ± 0.0 (0.1–0.1) | 0.1 ± 0.0 (0.1–0.2) | 0.2 ± 0.2 (0.1–0.2) | |
0.4 | 0.4 ± 0.1 (0.4–0.4) | 0.4 ± 0.0 (0.4– 0.5) | 0.4 ± 0.0 (0.4–0.5) | |
0.3 | 0.2± 0.0 (0.2–0.3) | 0.2 ± 0.0 (0.2–0.3) | 0.3 ± 0.0 (0.2–0.3) | |
0.7 | 0.72 ± 0.1 (0.6–0.8) | 0.8 ± 0.1 (0.7–0.9) | 0.8 ± 0.1 (0.7–1.0) | |
0.6 | 0.6 ± 0.0 (0.6–0.6) | 0.6 ± 0.0 (0.5–0.7) | 0.7 ± 0.1 (0.6–0.8) | |
1.1 | 0.0 ± 0.1 (0.9–1.1) | 1.0 ± 0.1 (1.0–1.2) | 1.3 ± 0.2 (1.0–1.6) | |
0.9 | 1.1 ± 0.1 (1.1–1.3) | 1.0 ± 0.2 (0.8–1.5) | 0.8 ± 0.1 (0.5–1.2) | |
0.8 | 0.8 ± 0.1 (0.6–0.8) | 0.7 ± 0.1 (0.6–0.9) | 0.7 ± 0.1 (0.6–0.8) | |
0.3 | 0.5 ± 0.1 (0.4–0.6) | 0.4 ± 0.1 (0.3–0.5) | 0.3 ± 0.1 (0.2–0.4) | |
0.8 | 0.9 ± 0.0 (0.9–1.0) | 0.8 ± 0.0 (0.7–0.9) | 0.8 ± 0.1 (0.7– 0.9) | |
0.9 | 0.8 ± 0.0 (0.8–0.9) | 0.8 ± 0.0 (0.7–0.9) | 1.0 ± 0.0 (0.9–1.1) | |
1.1 | 1.0 ± 0.0 (1.0–1.1) | 1.0 ± 0.0 (1.0–1.1) | 1.1 ± 0.1 (0.9–1.2) | |
0.1 | 0.1 ± 0.0 (0.1–0.1) | 0.1 ± 0.0 (0.1–1.1) | 0.2 ± 0.0 (0.9–0.2) |
Morphological measurements (mm) of adult female specimens of
|
|
|
|
|
Indonesia and Malaysia | (previously |
(previously |
||
Thailand | Thailand | |||
|
73.3 | 93.9 ± 7.0 | 95.5 ± 3.5 | 69.0 ± 10.1 |
(93.0–98.0) | (107.1–85.1) | (50.0–81.8) | ||
|
29.8 | 35.3 ± 1.1 | 37.5 ± 2.0 | 27.1 ± 4.0 |
(34.5–36.1) | (34.2–41.6) | (19.2–33.0) | ||
|
27.3 | 28.0 ± 1.1 | 36.2 ± 2.3 | 24.5 ± 4.1 |
(27.2–28.7) | (31.8–39.4) | (17.2–30.9) | ||
|
22.4 | 35.1 ± 1.1 | 28.1 ± 1.4 | 20.1 ± 2.8 |
(34.3–35.9) | (25.5–29.9) | (14.6–23.6) | ||
|
5.3 | 5.9 ± 0.3 | 6.9 ± 0.6 | 4.9 ± 0.7 |
(5.7–6.1) | (5.8–8.1) | (3.5–6.1) | ||
|
11.4 | 15.2 ± 0.3 | 16.3 ± 1.0 | 11.1 ± 1.6 |
(14.9–15.4) | (14.8–18.3) | (7.9–13.2) | ||
|
17.1 | 22.1 ± 0.8 | 21.5 ± 1.4 | 15.6 ± 2.2 |
(21.5–22.6) | (19.0–24.1) | (11.3–18.2) | ||
|
6.2 | 8.6 ± 0.3 | 9.0 ± 0.7 | 6.0 ± 0.8 |
(8.4–8.8) | (7.9–10.4) | (4.3–7.2) | ||
|
3.8 | 5.2 ± 0.9 | 4.5 ± 0.8 | 3.0 ± 0.8 |
(4.6–5.9) | (3.6–6.1) | (1.4–4.0) | ||
|
5.8 | 6.4 ± 0.6 | 6.5 ± 0.6 | 4.9 ± 0.6 |
(5.6–6.8) | (5.7–7.6) | (3.8–5.7) | ||
|
3.3 | 4.1 ± 0.2 | 4.7 ± 0.6 | 3.2 ± 0.5 |
(4.0–4.2) | (3.9–5.7) | (2.2–4.0) | ||
|
6.7 | 7.5 ± 0.2 | 8.5 ± 0.7 | 6.9 ± 0.7 |
(7.4–7.6) | (7.7–9.7) | (5.4–8.4) | ||
|
3.1 | 4.4 ± 0.5 | 4.8 ± 0.5 | 3.9 ± 0.64 |
(4.0–4.8) | (4.0–5.7) | (2.9–5.4) | ||
|
5.9 | 7.4 ± 0.4 | 7.5 ± 0.7 | 5.4 ± 0.8 |
(7.1–7.7) | (6.3–8.4) | (4.2–6.9) | ||
|
17.0 | 22.5 ± 1.5 | 22.5 ± 1.9 | 16.5 ± 2.3 |
(23.4–21.5) | (19.6–26.1) | (12.0–19.6) | ||
|
12.9 | 17.6 ± 0.3 | 17.6 ± 1.4 | 13.0 ± 2.2 |
(17.4–17.8) | (15.4–20.1) | (9.2–16.4) | ||
|
34.4 | 40.5 ± 4.6 | 43.8 ± 2.8 | 29.9 ± 4.3 |
(37.2–43.7) | (40.0–48.8) | (21.6–35.8) | ||
|
35.7 | 46.4 ± 0.5 | 48.0 ± 4.1 | 31.9 ± 4.5 |
(46.0–46.7) | (42.6–54.9) | (23.1–37.0) | ||
|
40.2 | 48.5 ± 0.7 | 49.2 ± 3.9 | 34.8 ± 5.0 |
(48.0–49.0) | (43.6–547) | (24.6–42.7) | ||
|
57.2 | 73.5 ± 1.7 | 73.1 ± 7.7 | 49.9 ± 7.0 |
(72.3–74.7) | (64.2–86.2) | (35.4–58.8) | ||
|
13.6 | 18.2 ± 0.6 | 17.8 ± 1.4 | 13.0 ± 2.1 |
(17.7–18.6) | (15.2–20.3) | (9.2–15.8) | ||
|
4.4 | 4.9 ± 0.5 | 5.7 ± 0.5 | 4.1 ± 0.7 |
(4.6–5.3) | (4.9–6.4) | (2.9–5.4) | ||
|
14.8 | 16.7 ± 0.2 | 17.5 ± 1.9 | 12.4 ± 1.8 |
(16.5–16.9) | (13.8–20.5) | (8.5–15.2) |