Research Article |
Siriporn Yodthong‡, Bryan L. Stuart§, Anchalee Aowphol‡
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Corresponding author: Anchalee Aowphol ( fsciacl@ku.ac.th ) Academic editor: Angelica Crottini
© 2019 Siriporn Yodthong, Bryan L. Stuart, Anchalee Aowphol.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Yodthong S, Stuart BL, Aowphol A (2019) Species delimitation of crab-eating frogs (Fejervarya cancrivora complex) clarifies taxonomy and geographic distributions in mainland Southeast Asia. ZooKeys 883: 119-153. https://doi.org/10.3897/zookeys.883.37544
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Abstract
The taxonomy and geographic distributions of species of crab-eating frogs (Fejervarya cancrivora complex) in mainland Southeast Asia have been highly uncertain. Three taxonomic names are used in recent literature (F. cancrivora, F. raja, and F. moodiei) but the applications of these names to localities has been inconsistent, especially owing to the lack of available molecular data for F. raja. Morphometric and mitochondrial DNA variation was examined in these frogs, including name-bearing types and topotypes of all three species. Findings corroborate evidence for the existence of two species in coastal mainland Southeast Asia, with F. moodiei having a wide geographic distribution and F. cancrivora sensu stricto occurring only in extreme southern Thailand and peninsular Malaysia. Fejervarya raja is shown to be only a large-bodied population of F. cancrivora sensu stricto and is synonymized with that species. Revised descriptions of F. moodiei and F. cancrivora sensu stricto are provided.
Keywords
Amphibia, cryptic species, Dicroglossidae, systematics, taxonomy
Introduction
Southeast Asia harbors high levels of amphibian species diversity and endemism (
Species of frogs in the genus Fejervarya Bolkay, 1915 have been subject to numerous investigations into cryptic diversity in efforts to resolve species boundaries and uncertain taxonomy in South, Southeast and East Asia (e.g.,
Two of these species, F. cancrivora and F. raja, have been reported from Thailand, where they occur in the vicinity of sea shores or river mouths (
In this study, we examined morphology and mitochondrial DNA variation in historical and newly-collected museum specimens of the F. cancrivora complex from Thailand and adjacent Asian countries to evaluate and clarify the taxonomic status of F. cancrivora, F. moodiei and F. raja. Importantly, our analyses included molecular and morphological data of topotypes of F. raja, and morphological data from the name-bearing type specimens of F. cancrivora and F. moodiei.
Materials and methods
Sampling
During 2015–2017, specimens of F. cancrivora were collected at 12 localities and F. raja at two localities in Thailand (Fig.
Map of sampling localities of the Fejervarya cancrivora complex, including F. cancrivora neotype (yellow pentagon), F. cancrivora sensu stricto (yellow circles), F. moodiei holotype (blue diamond), F. moodiei (blue triangles), and F. cancrivora samples that were referred to F. raja (red circles) prior to this study. Open symbols indicate molecular data only, shaded symbols indicate morphological data only, and shaded symbols with center dots indicate both molecular and morphological data were studied.
Specimens of Fejervarya used in (A) molecular and/or (B) morphological analyses.
| Species identification | Locality | Museum No. | GenBank Acession No. | Type of analyses | Reference | |
|---|---|---|---|---|---|---|
| Previous study | This study | |||||
| F. moodiei (holotype) | F. moodiei | Manila, Luzon, Philippines | CM 3724 | – | B | This study |
| F. cancrivora | F. moodiei | Malaysia | CNHM 161312 | – | B | This study |
| F. cancrivora | F. moodiei | Northern Luzon | FMNH 161693 | – | B | This study |
| F. cancrivora | F. moodiei | Northern Luzon | FMNH 161697 | – | B | This study |
| F. cancrivora | F. moodiei | Chonburi, Thailand | FMNH 190532 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Surat Thani, Surat Thani, Thailand | THNHM 05857 | – | B | This study |
| F. cancrivora | F. moodiei | Moo Ko Chumphon National Park, Chumphon, Thailand | THNHM 01032 | – | B | This study |
| F. cancrivora | F. moodiei | Moo Ko Chumphon National Park, Chumphon, Thailand | THNHM 01031 | – | B | This study |
| F. cancrivora | F. moodiei | Moo Ko Chumphon National Park, Chumphon, Thailand | THNHM 01033 | – | B | This study |
| F. cancrivora | F. moodiei | Ko Libong, Trang, Thailand | THNHM 02249 | – | B | This study |
| F. cancrivora | F. moodiei | Songkhla lake, Songkhla, Thailand | THNHM 02405 | – | B | This study |
| F. cancrivora | F. moodiei | Songkhla lake, Phatthalung, Thailand | THNHM 04332 | – | B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | THNHM 14252 | – | B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | THNHM 14254 | – | B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | THNHM 14255 | – | B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | THNHM 14256 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Trat, Trat, Thailand | THNHM 16631 | – | B | This study |
| F. cancrivora | F. moodiei | Tak Bai, Narathiwat, Thailand | THNHM 19720 | – | B | This study |
| F. cancrivora | F. moodiei | Tak Bai, Narathiwat, Thailand | THNHM 19721 | – | B | This study |
| F. cancrivora | F. moodiei | Tak Bai, Narathiwat, Thailand | THNHM 19724 | – | B | This study |
| F. cancrivora | F. moodiei | Tak Bai, Narathiwat, Thailand | THNHM 19725 | – | B | This study |
| F. cancrivora | F. moodiei | Suk Samran, Ranong, Thailand | THNHM 25736 | – | B | This study |
| F. cancrivora | F. moodiei | Suk Samran, Ranong, Thailand | THNHM 26002 | – | B | This study |
| F. cancrivora | F. moodiei | Suk Samran, Ranong, Thailand | THNHM 26016 | – | B | This study |
| F. cancrivora | F. moodiei | Sam Roi Yot, Prachuap Khiri Khan, Thailand | ZMKU AM 01368 | MN453492 | A | This study |
| F. cancrivora | F. moodiei | Sam Roi Yot, Prachuap Khiri Khan, Thailand | ZMKU AM 01369 | MN453493 | A, B | This study |
| F. cancrivora | F. moodiei | Sam Roi Yot, Prachuap Khiri Khan, Thailand | ZMKU AM 01370 | MN453494 | A | This study |
| F. cancrivora | F. moodiei | Sam Roi Yot, Prachuap Khiri Khan, Thailand | ZMKU AM 01371 | – | B | This study |
| F. cancrivora | F. moodiei | Kraburi, Ranong, Thailand | ZMKU AM 01373 | MN453495 | A, B | This study |
| F. cancrivora | F. moodiei | Kraburi, Ranong, Thailand | ZMKU AM 01375 | MN453496 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang, Phuket, Thailand | ZMKU AM 01376 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang, Phuket, Thailand | ZMKU AM 01377 | MN453497 | A | This study |
| F. cancrivora | F. moodiei | Mueang, Phuket, Thailand | ZMKU AM 01381 | MN453498 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Samui, Surat Thani, Thailand | ZMKU AM 01384 | MN453499 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Samui, Surat Thani, Thailand | ZMKU AM 01386 | – | B | This study |
| F. cancrivora | F. moodiei | Ko Samui, Surat Thani, Thailand | ZMKU AM 01387 | MN453500 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang Phang-nga, Phang-nga, Thailand | ZMKU AM 01390 | MN453501 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang Phang-nga, Phang-nga, Thailand | ZMKU AM 01394 | MN453502 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang Phang-nga, Phang-nga, Thailand | ZMKU AM 01397 | MN453503 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang Phang-nga, Phang-nga, Thailand | ZMKU AM 01398 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Phuket, Phuket, Thailand | ZMKU AM 01399 | MN453504 | A, B | This study |
| F. cancrivora | F. moodiei | Mueang Phuket, Phuket, Thailand | ZMKU AM 01400 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Phuket, Phuket, Thailand | ZMKU AM 01404 | – | B | This study |
| F. cancrivora | F. moodiei | Ko Lanta, Krabi, Thailand | ZMKU AM 01405 | MN453505 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Lanta, Krabi, Thailand | ZMKU AM 01407 | – | B | This study |
| F. cancrivora | F. moodiei | Ko Lanta, Krabi, Thailand | ZMKU AM 01409 | MN453506 | A | This study |
| F. cancrivora | F. moodiei | Ko Lanta, Krabi, Thailand | ZMKU AM 01413 | MN453507 | A | This study |
| F. cancrivora | F. moodiei | Khanom, Nakhon Si Thammarat, Thailand | ZMKU AM 01436 | – | B | This study |
| F. cancrivora | F. moodiei | Ko Chang, Trat, Thailand | ZMKU AM 01442 | MN453508 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Chang, Trat, Thailand | ZMKU AM 01446 | MN453509 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Chang, Trat, Thailand | ZMKU AM 01451 | MN453510 | A, B | This study |
| F. cancrivora | F. moodiei | Ko Chang, Trat, Thailand | ZMKU AM 01453 | – | B | This study |
| F. cancrivora | F. moodiei | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01467 | MN453511 | A, B | This study |
| F. cancrivora | F. moodiei | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01469 | – | B | This study |
| F. cancrivora | F. moodiei | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01470 | – | B | This study |
| F. cancrivora | F. moodiei | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01475 | MN453512 | A, B | This study |
| F. cancrivora | F. moodiei | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01479 | MN453513 | A, B | This study |
| F. cancrivora | F. moodiei | Kraburi, Ranong, Thailand | ZMKU AM 01485 | MN453514 | A, B | This study |
| F. cancrivora | F. moodiei | Kraburi, Ranong, Thailand | ZMKU AM 01486 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Krabi, Krabi, Thailand | ZMKU AM 01488 | – | B | This study |
| F. cancrivora | F. moodiei | Mueang Krabi, Krabi, Thailand | ZMKU AM 01489 | – | B | This study |
| F. cancrivora | F. moodiei | Kui Buri, Prachuap Khiri Khan, Thailand | ZMKU AM 01492 | – | B | This study |
| F. cancrivora | F. moodiei | La-ngu, Satun, Thailand | ZMKU AM 01493 | MN453515 | A, B | This study |
| F. cancrivora | F. moodiei | La-ngu, Satun, Thailand | ZMKU AM 01494 | – | B | This study |
| F. cancrivora | F. moodiei | La-ngu, Satun, Thailand | ZMKU AM 01498 | MN453516 | A, B | This study |
| F. cancrivora | F. moodiei | La-ngu, Satun, Thailand | ZMKU AM 01503 | MN453517 | A, B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | ZMKU AM 01516 | MN453518 | A, B | This study |
| F. cancrivora | F. moodiei | Kleang, Rayong, Thailand | ZMKU AM 01520 | MN453519 | A, B | This study |
| F. cancrivora | F. moodiei | Manila, Philippines | – | AB070738 | A |
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| F. cancrivora | F. moodiei | Negros Island, Philippines | – | AF206473 | A |
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| F. cancrivora | F. moodiei | Hainan, China | – | DQ458252 | A |
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| F. moodiei | F. moodiei | Dacope, Khulna, Bangladesh | – | AB530508 | A |
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| F. moodiei | F. moodiei | Teknaf, Cox’s Bazar, Bangladesh | – | AB543602 | A |
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| F. cancrivora | F. cancrivora | Cianjur, Java, Indonesia | – | AB444684 | A |
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| F. cancrivora | F. cancrivora | Padang, Sumatra, Indonesia | – | AB444685 | A |
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| F. cancrivora | F. cancrivora | Selangor, Malaysia | – | AB444688 | A |
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| F. cancrivora | F. cancrivora | Bogor, Java, Indonesia | – | AB444689 | A |
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| F. cancrivora | F. cancrivora | Banyumas, Java, Indonesia | – | AB444690 | A |
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| F. cancrivora | F. cancrivora | Malang, East Java, Indonesia | – | AB570273 | A |
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| F. cancrivora | F. cancrivora | Denpasar, Bali, Indonesia | – | AB570277 | A |
|
| F. cancrivora (neotype) | F. cancrivora | Cianjur, Java, Indonesia | FMNH 256688 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | CNHM 131093 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | CNHM 131100 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | CMNH 161102 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | CNHM 313095 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | FMNH 131108 | – | B | This study |
| F. cancrivora | F. cancrivora | Java, Indonesia | FMNH 131111 | – | B | This study |
| F. raja | F. cancrivora | Nakhon Si Thammarat, Thailand | FMNH 174052 | – | B | This study |
| F. raja | F. cancrivora | Phatthalung, Thailand | FMNH 174053 | – | B | This study |
| F. raja | F. cancrivora | Phatthalung, Thailand | FMNH 175923 | – | B | This study |
| F. raja | F. cancrivora | Phatthalung, Thailand | FMNH 175924 | – | B | This study |
| F. raja | F. cancrivora | Phatthalung, Thailand | FMNH 175925 | – | B | This study |
| F. raja | F. cancrivora | Phatthalung, Thailand | FMNH 175926 | – | B | This study |
| F. raja | F. cancrivora | Songkhla, Thailand | THNHM 04955 | – | B | This study |
| F. raja | F. cancrivora | Songkhla, Thailand | THNHM 04956 | – | B | This study |
| F. raja | F. cancrivora | Nong Chick, Pattani, Thailand | THNHM 15623 | – | B | This study |
| F. raja | F. cancrivora | Su-Ngai Kolok, Narathiwat, Thailand | THNHM 19221 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19771 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19765 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19766 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19767 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19768 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19769 | – | B | This study |
| F. raja | F. cancrivora | Tak Bai, Narathiwat, Thailand | THNHM 19770 | – | B | This study |
| F. raja | F. cancrivora | Pak Phayun, Phatthalung, Thailand | THNHM 19852 | – | B | This study |
| F. raja | F. cancrivora | Pak Phayun, Phatthalung, Thailand | THNHM 19853 | – | B | This study |
| F. raja | F. cancrivora | Pak Phayun, Phatthalung, Thailand | THNHM 19854 | – | B | This study |
| F. raja | F. cancrivora | Pak Phayun, Phatthalung, Thailand | THNHM 19855 | – | B | This study |
| F. raja | F. cancrivora | Pak Phayun, Phatthalung, Thailand | THNHM 19857 | – | B | This study |
| F. raja | F. cancrivora | Su-Ngai Kolok, Narathiwat, Thailand | THNHM 20754 | – | B | This study |
| F. raja | F. cancrivora | Nong Chick, Pattani, Thailand | THNHM 21248 | – | B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | THNHM 25499 | – | B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01418 | MN453520 | A | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01423 | MN453521 | A, B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01424 | – | B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01425 | MN453522 | A | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01426 | MN453523 | A, B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01429 | – | B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01430 | MN453524 | A, B | This study |
| F. raja | F. cancrivora | Khuan Khanun, Phatthalung, Thailand | ZMKU AM 01432 | – | B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01507 | MN453525 | A, B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01508 | – | B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01509 | MN453526 | A, B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01510 | – | B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01511 | MN453527 | A, B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01512 | – | B | This study |
| F. raja | F. cancrivora | Pak Phanang, Nakhon Si Thammarat, Thailand | ZMKU AM 01513 | – | B | This study |
| Fejervarya sp. | Fejervarya sp. | Pelabuhan ratu, Java, Indonesia | – | AB444693 | A |
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| Fejervarya sp. | Fejervarya sp. | Makassar, Sulawesi, Indonesia | – | AB570278 | A |
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| Fejervarya sp. | Fejervarya sp. | Makassar, Sulawesi, Indonesia | – | AB570288 | A |
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| F. cancrivora | Fejervarya sp. | Selatan, Sulawesi, Indonesia | – | EU979849 | A |
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| F. iskandari | F. iskandari | Malang, Java, Indonesia | – | AB570268 | A |
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| F. limnocharis | F. limnocharis | Java, Indonesia | – | AB277292 | A |
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| F. multistriata | F. multistriata | Yunan, China | – | AB354237 | A |
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| F. vittigera | F. vittigera | Quezon, Luzon Island, Philippines | – | AY313683 | A |
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| Euphlyctis cyanophlyctis | E. cyanophlyctis | Mangalore, India | – | AB488901 | A |
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| Limnonectes jarujini | L. jarujini | Surat Thani, Thailand | – | AB558951 | A |
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| Occidozyga lima | O. lima | Kuala Lumpur, Malaysia | – | AB488903 | A |
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DNA extraction, amplification and sequencing
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 (
Phylogenetic analysis
Homologous sequences of F. cancrivora and F. moodiei, and the outgroup taxa F. iskandari Vieth, Kosuch, Ohler & Dubois, 2001, F. limnocharis (Gravenhorst, 1829), F. multistriata (Hallowell, 1861), F. vittigera (Wiegmann, 1834), Euphlyctis cyanophlyctis (Schneider, 1799), Limnonectes jarujini Matsui, Panha, Khonsue & Kuraishi, 2010, and Occidozyga lima (Gravenhorst, 1829) (following
Morphological study
Morphological analyses were performed on 108 sexually mature individuals (61 males, 47 females) of F. cancrivora, F. moodiei, and F. raja (Table
Measurements were taken with digital Vernier calipers to the nearest 0.1 mm. Twenty-three morphological characters were measured following
EL eye length, greatest diameter of the eye including upper eyelids,
EN distance from front of eye to nostril,
FAL forelimb length, from elbow to base of outer palmar tubercle,
FOL foot length, from base of inner metatarsal tubercle to tip of fourth toe,
HAL hand length, from base of outer palmar tubercle to tip of third finger,
HL head length, from back of mandible to tip of snout,
HLL hindlimb length,
HW head width, from left side back of mandible to right side back of mandible,
IMTL length of inner metatarsal tubercle,
IN internarial space, distance between the nostrils,
IOD interorbital distance,
ITL inner toe length,
NS nostril-snout length, distance from nostril to tip of snout,
NTL nostril-tympanum length, distance between nostril and front of tympanum,
SL snout length, distance from front of eye to tip of snout,
STL snout-tympanum length, tip of snout to front of tympanum,
SVL snout-vent length,
TD tympanum diameter, maximum diameter,
TEL tympanum-eye length, distance between end of eye to front of tympanum,
TFOL length of tarsus and foot, from base of tarsus to tip of fourth toe,
THIGHL thigh length,
TL tibia length,
UEW maximum width of upper eyelids,
1FL first finger length.
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 SVL to a ratio (r) and then converted to a percentage. Specimens were assigned to group (= species) based on their mtDNA assignment (below). Principal component analysis (PCA) was performed separately by sex using FactoMineR and factoextra R package (
Results
Phylogenetic analyses
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 F. cancrivora complex as monophyletic with strong support, and to contain two major clades referred to as Clades A and B (Fig.
Uncorrected pairwise sequence divergences (p-distances) were relatively low within subclades, with subclade A1 ranging from 0.6–6.0% (mean 3.6%), subclade A2 ranging from 0.0–1.4% (mean 0.3%), and subclades B1 and B2 each ranging from 0.0–1.6% (means 0.4%; Table
Bayesian consensus phylogram of the mitochondrial16S rRNA gene of Fejervarya cancrivora and the closely related species, F. moodiei and F. raja. Numbers at nodes represent Bayesian posterior probability support values. Clade and subclade names are presented next to branches and group names are presented to the right of terminal taxa.
Uncorrected pairwise sequence divergences (p-distances) in the mitochondrial 16S rRNA gene of Fejervarya cancrivora and related species. Mitochondrial subclades A1, A2, B1, and B2 are defined in the text.
| iskandari | multistriata | limnocharis | vittigera | cancrivora B2 | cancrivora B1 | cancrivora A2 | sp. A1 | |
| iskandari | – | |||||||
| multistriata | (12.8) | – | ||||||
| 12.8 | ||||||||
| limnocharis | (12.1–12.7) | (0.2–0.4) | (0.9) | |||||
| 12.4 | 0.3 | 0.9 | ||||||
| vittigera | (16.2) | (12.2) | (11.7–13.5) | – | ||||
| 16.2 | 12.2 | 12.6 | ||||||
| cancrivora B2 | (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 | ||||
| cancrivora B1 | (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 | |||
| cancrivora A2 | (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 | ||
| sp. A1 | (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 |
Morphological analyses
PCA analysis of males revealed morphometric differences between F. cancrivora Group A and F. cancrivora Group B, with no overlap on a plot of the first two axes (Fig.
PCA analysis of females revealed morphometric differences between F. cancrivora Group A and F. cancrivora Group B, with only slight overlap on a plot of the first two axes (Fig.
Factor loading on the first three principal components of 23 morphological characters for male and female Fejervarya cancrivora, F. moodiei, and F. raja.
| Character | Males | Females | ||||
|---|---|---|---|---|---|---|
| PC 1 | PC 2 | PC 3 | PC 1 | PC 2 | PC 3 | |
| SVL | 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 Fejervarya cancrivora and F. moodiei. Data are given as mean and standard deviation, followed by range in parentheses. Key: a tested by Mann-Whitney U test, * significance level at p < 0.05.
| Characters | Males | Females | ||||||
|---|---|---|---|---|---|---|---|---|
| F. cancrivora | F. moodiei | t-test | p | F. cancrivora | F. moodiei | t-test | p | |
| n = 31 | n = 30 | n = 14 | n = 33 | |||||
| SVL | 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) | |||||
| HL/HW | 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) | |||||
| IOD/HW | 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) | |||||
| SL/HL | 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) | |||||
| EL/HL | 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) | |||||
| NS/EN | 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) | |||||
| EL/SL | 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) | |||||
| EL/EN | 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) | |||||
| IN/IOD | 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) | |||||
| TD/EL | 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) | |||||
| TEL/EL | 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) | |||||
| FAL/HAL | 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) | |||||
| THIGHL/TL | 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) | |||||
| FOL/TL | 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) | |||||
| IMTL/TL | 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) | |||||
Species accounts
The genetic and morphometric data provide congruent, independent lines of evidence to support the hypothesis that F. cancrivora Groups A and B represent two separate species. Specifically, Group A consists of a composite of F. cancrivora from Indonesia and Malaysia, and F. raja from Thailand (
Fejervarya cancrivora
Rana cancrivora
Gravenhorst, 1829: 41;
Rana cancrivora raja Smith, 1930: 96
Rana raja Taylor, 1962: 373; Stuart et al. 2006: 19
Fejervarya cancrivora: Dubois & Ohler, 2000: 35;
Fejervarya cancrivora: Large type
Fejervarya raja:
Diagnosis
Fejervarya cancrivora can be characterized by the following combination of characters: (1) large size, SVL 60.2–79.8 mm in males, 85.1–107.1 mm in females (Table
Description of neotype
Dubios and Ohler (2000) designated and described the neotype adult male, FMNH 256688, from Java, Indonesia (Fig.
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 A adult male neotype of Fejervarya cancrivora (FMNH 256688) B adult male F. cancrivora (ZMKU AM 01426) from Khuan Khanun District, Phatthalung Province, Thailand C adult female holotype of F. moodiei holotype (CM 3724), and D adult male F. moodiei (ZMKU AM 10390) from Mueang Phang-nga District, Phang-nga Province, Thailand. The inner metatarsal ridge on the tarsus of F. cancrivora is indicated with an arrow.
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.
Coloration of neotype in preservative
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.
Coloration of referred Thai specimen in life
Adult male ZMKU AM 01426 (Fig.
Variations
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 (n = 18, 41%) or absent (n = 26, 59%). Two specimens from Nakhon Si Thammarat Province, Thailand (ZMKU AM 01509 and ZMKU AM 01513), have a narrow light brown stripe on tibia. Pineal body not visible in one male specimen from Pattani Province, Thailand (THNHM 21248).
Distribution
Based on a combination of the morphological and genetic studies of F. cancrivora large type (
Habitat, ecology and natural history
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 A Fejervarya cancrivora at a wetland in Khuan Khanun District, Patthalung Province B F. cancrivora at a brackish shrimp pond near Pak Phanang river, Pak Phanang District, Nakhon Si Thammarat Province C F. moodiei at mangrove forest in Thai Mueang Distrinct, Phang-nga Province, and D F. moodiei at brackish fish ponds near mangroves at the mouth of the Prasae River, Kleang District, Rayong Province. Photograph A by Attapol Rujirawan.
Fejervarya moodiei
Rana moodiei Taylor, 1920: 234
Rana cancrivora:
Fejervarya moodiei:
Fejervarya cancrivora:
Fejervarya
Bangladesh mangrove type
Fejervarya cancrivora
mangrove type
Fejervarya cf. cancrivora Harikrishnan & Vasudevan, 2018: 241
Diagnosis
Fejervarya moodiei can be characterized by the following combination of characters: (1) medium to large size, SVL 42.7–62.7 mm in males, 50.0–81.8 mm in females (Table
Description of holotype
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 of holotype in preservative
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.
Coloration of referred Thai specimen in life
Adult male ZMKU AM 01390 (Fig.
Variations
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 (ZMKU AM 01444 and 01451) have dark orange markings on anterior part of dorsum. Markings or transverse spots on dorsum and dorsal surfaces of forelimbs and hindlimbs usually distinct, but faint in a few individuals. Coloration on flank usually creamy white, but pale brown, with dark brown marbling, in some individuals. Ventral coloration usually creamy white, but pale brown with indistinct dark mottling on chin and chest in some individuals. Hand usually creamy white, but light brown in some individuals. Most specimens have dermal fringe on fingers II and III (males N = 21, 70%; females N = 23, 69.7%), but some individuals lack this fringe (males N = 9, 30%; females N = 10, 30.3%). One specimen from Narathiwat Province, Thailand (THNHM 19720) has a vertebral stripe.
Distribution
Based on a combination of morphological and genetic studies of F. cancrivora mangrove type (
Habitat, ecology, and natural history
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.
Comparisons
Twelve species of Fejervarya are known (
Fejervarya cancrivora and F. moodiei differ from all of these species by having the following combination of characters: (1) medium to large body size (vs. small to medium, SVL about 30–40 mm in males for F. iskandari, F. kawamurai, F. limnocharis, SVL about 40–55 mm in males for F. multistriata, F. orissaensis, F. triora [
Fejervarya moodiei differs from F. cancrivora by having: (1) SVL 42.7–62.7 mm in males, 50.0–81.8 mm in females (vs. 60.2–79.8 mm in males, 85.1–107.1 mm in females of F. cancrivora, Table
Discussion
Our study clarifies that two species of crab-eating frogs (Fejervarya cancrivora complex) occur in mainland Southeast Asia: F. moodiei in coastal regions throughout mainland Southeast Asia, with replacement by F. cancrivora sensu stricto in extreme southern Thailand (on the Gulf of Thailand coast) and peninsular Malaysia. These findings corroborate those of
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 F. cancrivora and F. moodiei in mainland Southeast Asia. Our sampling did not reveal F. cancrivora and F. moodiei to occur in sympatry, but did find the two species to occur in shrimp ponds that were separated by only approximately 4.5 air-km (or 5.2 km following the river course) along the Pak Phanang River in Pak Phanang District, Nakhon Si Thammarat Province, Thailand (Fig.
Acknowledgments
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 (FMNH), Sunchai Makchai (THNHM), and Stephen Rogers, Jennifer Sheridan and Kaylin Martin (CM) facilitated the study of specimens in their care. David S. McLeod provided helpful suggestions with the morphological study. Attapol Rujirawan assisted with statistical analyses and provided photographs. Attapol Rujirawan, Korkwan Termprayoon, Natee Ampai, and Piyawan Puanprapai assisted with field work. David S. McLeod and Guinevere O. U. Wogan improved the manuscript.
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Appendix 1
Specimens examined.
Fejervarya cancrivora (Gravenhorst, 1829): Indonesia, Java: CNHM 131093, 131100, 131105–09, 161102, 191098, 313095, FMNH 131108, 131111; West Java, Cianjur FMNH 256688 (neotype); Thailand, Nakhon Si Thammarat Province: FMNH 174052–53; Pak Phanang District: (8°17.454'N, 100°11.229'E) ZMKU AM 01507–13; THNHM 25499; Narathiwat Province, Su-Ngai Kolok District: THNHM 19221, 20754; Tak Bai District: THNHM 19722–23, 19726–28, 19764–71; Pattani Province, Nong Chick District: THNHM 15623, 21248–49; Phatthalung Province: CNHM 175923–26; Khuan Khanun District: (7°45.580'N, 100°9.446'E): ZMKU AM 01415–26, 01427–29, (7°44.127'N, 100°8.635E) ZMKU AM 01430–34; Pak Phayun District: THNHM 19852–57; Songkhla Province: THNHM 04332, 04955–56
Fejervarya moodiei (Taylor, 1920): Malaysia: CNHM 161312; Philippines, Northern Luzon: FMNH 161693, 161697; Luzon, Manila: CM 3724 (holotype); Thailand: Chonburi Province: FNHM 190532, THNHM 04919–21, Mueang Chonburi District: THNHM 06408–12; Chumphon Province, Moo Ko Chumphon National Park: THNHM 01030–33; Krabi Province, Ko Lanta District (7°35.702'N, 99°4.272'E): ZMKU AM 01405–14, Mueang Krabi District (8°4.502'N, 98°55.506'E): ZMKU AM 01487–90; Nakhon Si Thammarat Province, Khanom District (9°12.760'N, 99°50.969'E): ZMKU AM 01435–41; Pak Phanang District (8°19.850'N, 100°11.870'E): ZMKU AM 01464–79; Narathiwat Province, Tak Bai District: THNHM 19720–21, 19724–25; Phang-nga Province, Mueang Phang-nga District (8°25.998'N, 98°30.973'E): ZMKU AM 01390–98; Phatthalung Province, Songkhla lake: THNHM 04332–33; Phuket Province, Mueang Phuket District (7°54.522'N, 98°24.425'E): ZMKU AM 01376–83, 01399–404; Prachuap Khiri Khan Province, Kui Buri District (12°8.143'N, 99°57.737'E): ZMKU AM 01491–92, Sam Roi Yot District: ZMKU AM 01368–71; Ranong Province, Kra Buri District (10°19.435'N, 98°45.894'E): ZMKU AM 01372–75, 01480–86, Suk Samran District: THNHM 25736; 26002, 26016; Rayong Province, Klaeng District THNHM 14252–64, (12°42.164'N, 101°41.634'E): ZMKU AM 01514–20; Samut Prakarn Province, Phra Pradaeng District, Bang Krachao Sub-district: THNHM 26075–78; Satun Province, La-ngu District (6°51.861'N, 99°45.484'E): ZMKU AM 01493–506; Songkhla Province, Songkhla lake: THNHM 02403–05; Surat Thani Province, Ko Samui District (9°33.220'N, 100°3.327'E): ZMKU AM 01384–89; Mueang Surat Thani District, Makham Tia Sub-district: THNHM 05857–58; Trat Province, Ko Chang District (12°0.178'N, 102°22.639'E): ZMKU AM 01442–63; Klong Yai District: THNHM14292–94; Mueang Trat District: THNHM 16631–36, 24452; Trang Province, Kantang District, Ko Libong: THNHM 02249.
Appendix 2
Morphological measurements (mm) of adult male specimens of Fejervarya. Data are given as mean and standard deviation, followed by range in parentheses.
| Characters | F. cancrivora neotype | F. cancrivora Indonesia and Malaysia | F. cancrivora (previously F. raja) Thailand | F. moodiei (previously F. cancrivora) Thailand |
|---|---|---|---|---|
| N = 1 | N = 4 | N = 26 | N = 30 | |
| SVL | 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) |
| HL | 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) |
| HW | 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) |
| STL | 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) |
| NS | 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) |
| SL | 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) |
| NTL | 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) |
| EN | 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) |
| TEL | 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) |
| TD | 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) |
| IN | 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) |
| EL | 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) |
| IOD | 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) |
| UEW | 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) |
| HAL | 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) |
| FAL | 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) |
| THIGHL | 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) |
| TL | 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) |
| FOL | 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) |
| TFOL | 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) |
| 1FL | 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) |
| IMTL | 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) |
| ITL | 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) |
| HL/HW | 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) |
| IOD/HW | 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) |
| SL/HL | 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) |
| EL/HL | 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) |
| NS/EN | 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) |
| EL/SL | 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) |
| EL/EN | 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) |
| IN/IOD | 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) |
| TD/EL | 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) |
| TEL/EL | 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) |
| FAL/HAL | 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) |
| THIGHL/TL | 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) |
| FOL/TL | 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) |
| IMTL/TL | 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) |
Appendix 3
Morphological measurements (mm) of adult female specimens of Fejervarya. Data are given as mean and standard deviation, followed by range in parentheses.
| Character | F. moodiei | F. cancrivora | F. cancrivora | F. moodiei |
| CM 3724 Holotype | Indonesia and Malaysia | (previously F. raja) | (previously F. cancrivora) | |
| Thailand | Thailand | |||
| N = 1 | N = 2 | N = 12 | N = 32 | |
| SVL | 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) | ||
| HL | 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) | ||
| HW | 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) | ||
| STL | 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) | ||
| NS | 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) | ||
| SL | 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) | ||
| NTL | 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) | ||
| EN | 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) | ||
| TEL | 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) | ||
| TD | 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) | ||
| IN | 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) | ||
| EL | 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) | ||
| IOD | 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) | ||
| UEW | 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) | ||
| HAL | 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) | ||
| FAL | 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) | ||
| THIGHL | 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) | ||
| TL | 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) | ||
| FOL | 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) | ||
| TFOL | 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) | ||
| 1FL | 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) | ||
| IMTL | 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) | ||
| ITL | 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) |