Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5785 No. 1: 27 Mar. 2026
Type: Article
Published: 2026-03-27
DOI: 10.11646/zootaxa.5785.1.3
Page range: 67-102
Abstract views: 81
PDF downloaded: 1

A systematic review of the Limnonectes hascheanus-limborgi complex (Anura: Dicroglossidae), with the description of a new species from Peninsular Malaysia

School of Biological Sciences; Universiti Sains Malaysia; 11800 USM; Penang; Malaysia.
School of Biological Sciences; Universiti Sains Malaysia; 11800 USM; Penang; Malaysia.; Centre for Marine and Coastal Studies; Universiti Sains Malaysia; 11800 USM; Penang; Malaysia.
School of Biological Sciences; Universiti Sains Malaysia; 11800 USM; Penang; Malaysia.
School of Biological Sciences; Universiti Sains Malaysia; 11800 USM; Penang; Malaysia.
Herpetology Laboratory; Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California; 92515; USA.
Herpetology Laboratory; Department of Biology; La Sierra University; 4500 Riverwalk Parkway; Riverside; California; 92515; USA.; Department of Herpetology; San Diego Natural History Museum; PO Box 121390; San Diego; California; 92112; USA.; El Serpentario y C.E.M.A de Baja California Sur; Brercha California e/ Nuevo Reforma y Guaycura. Fracc: Benito Juárez; CP 23090; La Paz; Baja California Sur; México
Institute for Tropical Biology and Conservation; Universiti Malaysia Sabah; Jalan UMS; 88400; Kota Kinabalu; Sabah; Malaysia.; Lee Kong Chian Natural History Museum; National University of Singapore; 2 Conservatory Drive; Singapore 117377
Amphibia Anuran Amphibian Frog cryptic species integrative taxonomy biogeography Southeast Asia

Abstract

The taxonomic confusion of the Limnonectes hascheanus-limborgi complex in Peninsular Malaysia has lasted for more than a decade since the presence of L. limborgi was reported in the country based on molecular data in 2014. Before that, L. hascheanus was thought to be the only species of L. hascheaus-limborgi complex occurring in Peninsular Malaysia. Currently, comprehensive approaches using molecular and morphological analyses corroborate the existence of a new species, Limnonectes corvus sp. nov. from Peninsular Malaysia. Limnonectes corvus sp. nov. is syntopic with L. hascheanus in some localities but differs by having an average uncorrected p-distance of 7.8% based on the 16S rRNA gene and the following diagnostic morphological characteristics: (1) small size with SVL 21.59–35.45 mm in males, SVL 28.74–35.78 mm in females; (2) short, stocky and robust body; (3) males lack external vocal sacs; (4) loreal region slightly concave; (5) tympanum approximately 2/3 of the eye diameter; (6) distinct supratympanic fold along the tympanum extending to supra-axillary region; (7) smooth skin without dense cluster of granular warts on super-axillary and cloacal region (8) fingertips with terminal bulbs; (9) relative finger lengths III>I=II>IV or III>IV=I=II; (10) inner metatarsal tubercle distinctly raised and less than 50% of the length of the first toe; (11) dermal ridge along the outer metatarsal; (12) toes basally webbed, with webbing extending to middle subarticular tubercle of the fourth toe with webbing formula as follow: I 1-–2 II 0+–22/3 III 1+–3 IV 3–11/2 V or I 1-–2 II 1+–2½ III 1½–3 IV 3–2- IV; (13) enlarged odontoids are present in both males and females. The current study found that previously reported populations of L. limborgi in Peninsular Malaysia are misidentifications of the new species L. corvus sp. nov. and we update its distribution to the east of the Bintang range to include the Sekayu lowland forest in Terengganu in the north and Ketumbar Hill, Kuala Lumpur, and Tioman Island in the south.

 

References

  1. AmphibiaWeb (2024) Information on amphibian biology and conservation. Available from: https://amphibiaweb.org (accessed 8 May 2024)
  2. Andersson, L.G. (1916) Zoological results of the Swedish zoological expeditions to Siam 1911–1912 and 1914. 3. Batrachians. Kungliga Svenska Vetenskaps-Akademiens Handlingar, 55, 13–17.
  3. Badli-Sham, B.H., Syafiq, M.F., Aziz, M.S.A., Mohd, J.N.R., Awang, M.T., Othman, M.N.A., Abdul A.A.A., Dzu, K., Abdol, W.N.A., Jamil, N.L., Ismail, M.A., Wahab, N.A.A., Xin, W.O., Jamaha, N.A.N., Aqmal-Naser, M., Fahmi-Ahmad, M, Shahirah-Ibrahim, N., Rizal, S.A., Belabut, D.M., Chan, K.O. & Ahmad, A.B. (2023) A decade of amphibian studies (Animalia, Amphibia) at Sekayu lowland forest, Hulu Terengganu, Peninsular Malaysia. ZooKeys, 1157, 43–93. https://doi.org/10.3897/zookeys.1157.95873
  4. Belabut, D.M. (2010) An Updated Checklist of the Herpetofauna of Pulau Singa Besar, Langkawi, Peninsular Malaysia. Malaysian Applied Biology, 39, 13–23.
  5. Berry, P.Y. (1975) The Amphibian Fauna of Peninsular Malaysia. Tropical Press, Kuala Lumpur, 110 pp.
  6. Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences, 97 (12), 6585–6590. https://doi.org/10.1073/pnas.97.12.6585
  7. Boulenger, G.A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Zoological Survey of India, 20, 1–223. https://doi.org/10.26515/rzsi/v20/i1/1920/163533
  8. Camargo, A. (2022) PCAtest: testing the statistical significance of Principal Component Analysis in R. PeerJ, 20, e12967. https://doi.org/10.7717/peerj.12967
  9. Chan-ard, T. (2003) A Photographic Guide to Amphibians in Thailand. Darnsutha Press Co., Ltd, Bangkok, Thailand, 175 pp. [in Thai]
  10. Chan, K.O., Abraham, R.K., Grismer, L.L. & Brown, R.M. (2020a) A systematic review of the Pulchrana picturata complex, with the description of a new species from Peninsular Malaysia, Sumatra, and southern Thailand. Raffles Bulletin of Zoology, 68, 880–890. https://doi.org/10.26107/rbz-2020-0096
  11. Chan, K.O., Abraham, R.K. & Badli-Sham, B.H. (2020b) A revision of the Asian tree toad complex Rentapia hosii (Anura: Bufonidae) with the description of a new species from Peninsular Malaysia. Raffles Bulletin of Zoology, 68, 595–607. https://doi.org/10.26107/RBZ-2020-0075
  12. Chan, K.O., Abraham, R.K., Grismer, J.L. & Grismer, L.L. (2018) Elevational size variation and two new species of torrent frogs from Peninsular Malaysia (Anura: Ranidae: Amolops Cope). Zootaxa, 4434 (2), 250–264. https://doi.org/10.11646/zootaxa.4434.2.2
  13. Chan, K.O., Alexander, A.M., Grismer, L.L., Su, Y.C., Grismer, J.L., Quah, E.S.H. & Brown, R.M. (2017) Species delimitation with gene flow: a methodological comparison and population genomics approach to elucidate cryptic species boundaries in Malaysian Torrent Frogs. Molecular Ecology, 26 (20), 5435–5450. https://doi.org/10.1111/mec.14296
  14. Chan, K.O. & Grismer, L.L. (2021) A standardized and statistically defensible framework for quantitative morphological analyses in taxonomic studies. Zootaxa, 5023 (2), 293–300. https://doi.org/10.11646/zootaxa.5023.2.9
  15. Chan, K.O. & Grismer, L.L. (2022) GroupStruct: An R package for allometric size correction. Zootaxa, 5124 (4), 471–482. https://doi.org/10.11646/zootaxa.5124.4.4
  16. Chan, K.O., Muin, M.A., Anuar, S., Andam, J., Razak, N. & Aziz, M.A. (2019) First checklist on the amphibians and reptiles of Mount Korbu, the second highest peak in Peninsular Malaysia. Check List, 15 (6), 1055–1069. https://doi.org/10.15560/15.6.1055
  17. Dubois, A. (1987) Miscellanea nomenclatorica batrachologica. Alytes, 3 (3), 7–95.
  18. Dasi, O. & Shahriza, S. (2020) A checklist of amphibians at Lubuk Semilang recreational park, Langkawi Island, Kedah, Peninsular Malaysia. Arxius de Miscellània Zoològica, 18, 9–26. https://doi.org/10.32800/amz.2020.18.0009
  19. Dehling, J.M. (2014) Eine neue Fangzahnfroschart der Gattung Limnonectes (Anura: Dicroglossidae) vom Gunung Lawit, Malaiische Halbinsel. Sauria, 36 (4), 17.
  20. Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7 (1), 214. https://doi.org/10.1186/1471-2148-7-214
  21. Emerson, S.B. & Berrigan, D. (1993) Systematics of Southeast Asian ranids: multiple origins of voicelessness in the subgenus Limnonectes (Fitzinger). Herpetologica, 49, 22–31.
  22. Fahmi, M., Kadafi, A.M., Priambodo, B., Fauzi, M.A., Hamidy, A., Firdaus, A.S., Smith, E.N. & Kurniawan, N. (2023) Phylogenetic Inference on Limnonectes kuhlii Complex in Java and Sumatra Reveals Significant Novel Diversity. The Journal of Experimental Life Science, 13 (3), 145–152. https://doi.org/10.21776/ub.jels.2023.013.03.02.
  23. Fei, L., Ye, C.Y. & Jiang, J.P. (2010) Phylogenetic systematics of Ranidae. Herpetologica Sinica, 12, 1–43.
  24. Flower, S.S. (1899) Notes on a second collection of batrachians made in the Malay peninsula and Siam, from November 1896 to September 1898, with a list of the species recorded from those countries. Proceedings of the Zoological Society of London, 1899, 885–916. https://doi.org/10.1111/j.1469-7998.1899.tb06880.x
  25. Frost, D.R. (2026) Amphibian Species of the World: an Online Reference. Version 6.2. American Museum of Natural History, New York, New York. Available from: https://amphibiansoftheworld.amnh.org/index.php (accessed 4 January 2026) https://doi.org/10.5531/db.vz.0001
  26. Garg, S., Chandrakasan, S., Gokulakrishnan, G., Gopika, C., Das, I. & Biju, S.D. (2022) The curious case of Charles Darwin’s frog, Rana charlesdarwini Das, 1998: Phylogenetic position and generic placement, with taxonomic insights on other minervaryan frogs (Dicroglossidae: Minervarya) in the Andaman and Nicobar Archipelago. Vertebrate Zoology, 72, 169–199. https://doi.org/10.3897/vz.72.e79496
  27. Gillespie, G., Ahmad, E., Scriven, S. & Shia, A. (2021) Field Guide to the Frogs of the Lower Kinabatangan Region, Sabah. Hutan (Kinabatangan Orang-utan Conservation Programme), Sabah, 71 pp.
  28. Gorin, V.A., Scherz, M.D., Korost, D.V. & Poyarkov, N.A. (2021) Consequences of parallel miniaturisation in Microhylinae (Anura, Microhylidae), with the description of a new genus of diminutive South East Asian frogs. Zoosystematics and Evolution, 97 (1), 21–54. https://doi.org/10.3897/zse.97.57968
  29. Grismer, J.L., Grismer, L.L., Das, I., Yaakob, N.S., Liat, L.B., Leong, Z.M., Youmans, T.M. & Kaiser, H.I. (2004a) Species diversity and checklist of the herpetofauna of Pulau Tioman, Peninsular Malaysia with a preliminary overview of habitat utilization. Asiatic Herpetological Research, 10, 247–279.
  30. Grismer, L.L. (2011) Field Guide to the Amphibians and Reptiles of the Seribuat Archipelago, Peninsular Malaysia. Edition Chimaira, Frankfurt am Main, 258 pp.
  31. Grismer, L.L., Chan, K.O., Grismer, J.L., Wood, P.L.J. & Norhayati, A. (2010) A checklist of the herpetofauna of the Banjaran Bintang, Peninsular Malaysia. Russian Journal of Herpetology, 17 (2), 147–160.
  32. Grismer, L.L., Rujirawan, A., Yodthong, S., Stuart, B.L., Le, M.D., Le, D.T., Chuaynkern, Y., Wood, P.L.J. & Aowphol, A. (2022) The taxonomy and phylogeny of the Cyrtodactylus brevipalmatus group (Squamata: Gekkonidae) with emphasis on C. interdigitalis and C. ngati. Vertebrate Zoology, 72, 245–269. https://doi.org/10.3897/vz.72.e80615
  33. Grismer, L.L., Sukumaran, J., Grismer, J.L., Youmans, T.M., Wood, P.L.J. & Johnson, R. (2004b) Report on the herpetofauna of the Temengor Forest Reserve, Perak, West Malaysia. Hamadryad, 29, 15–32.
  34. Grismer, L.L., Wood, P.L.J., Anuar, S., Quah, E.S.H., Muin, M.A., Chan, K.O., Sumarli, A.X. & Loredo, A.I. (2015) Repeated evolution of sympatric, paleoendemic species in closely related, co-distributed lineages of Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) across a sky-island archipelago in Peninsular Malaysia. Zoological Journal of the Linnean Society, 174 (4), 859–876. https://doi.org/10.1111/zoj.12254
  35. Grismer, L.L., Wood, P.L.J., Anuar, S., Muin, M.A., Quah, E.S.H., McGuire, J.A., Brown, R.M., Tri, V.N. & Hong Thai, P. (2013) Integrative taxonomy uncovers high levels of cryptic species diversity in Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) and the description of a new species from Peninsular Malaysia. Zoological Journal of the Linnean Society, 169 (4), 849–880. https://doi.org/10.1111/zoj.12064
  36. Grismer, L.L., Youmans, T.M., Wood, P.L.J., Ponce, A., Wright, S.B., Jones, B.S., Johnson, R., Sanders, K.L., Gower, D.J., Yaakob, N.S. & Lim, K.K. (2006) Checklist of the herpetofauna of Pulau Langkawi, Malaysia, with comments on taxonomy. Hamadryad, 30 (1/2), 61–74.
  37. Guayasamin, J.M., Bustamante, M.R., Almeida-Reinoso, D. & Funk, W.C. (2006) Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnean Society, 147 (4), 489–513. https://doi.org/10.1111/j.1096-3642.2006.00223.x
  38. Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59 (3), 307–321. https://doi.org/10.1093/sysbio/syq010
  39. Hamidi, J. (2013) Herpetofauna di Gunung Gagau, Terengganu. In: Latiff, A., Ali, C.A. & Mohamad, K.R. (Eds.), Gunung Gagau, Terengganu: Transforming Natural Assets into an Ecotourism Product. Academy of Sciences Malaysia and Ketengah, Kuala Lumpur, pp. 165–172.
  40. Harikrishnan, S. & Vasudevan, K. (2018) Amphibians of the Andaman & Nicobar Islands: distribution, natural history, and notes on taxonomy. Alytes, 36 (1–4), 238–265.
  41. Herlambang, A.E., Riyanto, A., Munir, M.A., Hamidy, A., Kimura, K., Eto, K. & Mumpuni, M. (2022) After 16 years: an update checklist of herpetofauna on the Natuna Islands, Indonesia. Treubia, 49 (2), 67–84. https://doi.org/10.14203/treubia.v49i2.4379
  42. Herr, M.W., Vallejos, J.G., Meneses, C.G., Abraham, R.K., Otterholt, R., Siler, C.D., Rico, E.L. & Brown, R.M. (2021) A new, morphologically cryptic species of fanged frog, genus Limnonectes (Amphibia: Anura: Dicroglossidae), from Mindoro Island, central Philippines. Ichthyology & Herpetology, 109 (1), 188–210. https://doi.org/10.1643/h2020095
  43. Hong, Z., Anuar, S., Grismer, L.L. & Quah, E.S.H. (2021) Preliminary report on the herpetofaunal diversity of Batu Hampar Recreational Forest, Kedah, Malaysia. Check List, 17 (3), 791–814. https://doi.org/10.15560/17.3.791
  44. Howlader, M.S., Nair, A. & Merilä, J. (2016) A new species of frog (Anura: Dicroglossidae) discovered from the mega city of Dhaka. PLoS One, 11 (3), e0149597. https://doi.org/10.1371/journal.pone.0149597
  45. Huang, J., Li, Z., Liu, X., Stuart, B.L., Yuan, Z. & Zhao, H. (2022) The phylogenetic status of Limnonectes liui (Yang, 1983) (Anura: Dicroglossidae) based on mitochondrial genes and its taxonomic implications. Zootaxa, 5092 (1), 116–126. https://doi.org/10.11646/zootaxa.5092.1.6
  46. Huelsenbeck, J.P. & Hillis, D.M. (1993) Success of phylogenetic methods in the four-taxon case. Systematic Biology, 42 (3), 247–264. https://doi.org/10.1093/sysbio/42.3.247
  47. Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17 (8), 754–755. https://doi.org/10.1093/bioinformatics/17.8.754
  48. Inger, R.F. & Stuart, B.L. (2010) Systematics of Limnonectes (Taylorana) Dubois. Current Herpetology, 29 (2), 51–68. https://doi.org/10.3105/018.029.0201
  49. Inger, R.F. & Stuebing, R.B. (2005) A Field Guide to the Frogs of Borneo. Second Edition. Natural History Publications (Borneo) Sdn Bhd, Kota Kinabalu, 201 pp.
  50. Jestrzemski, D., Schütz, S., Nguyen, T.Q. & Ziegler, T. (2013) A survey of amphibians and reptiles in Chu Mom Ray National Park, Vietnam, with implications for herpetofaunal conservation. Asian Journal of Conservation Biology, 2 (2), 88–110.
  51. Jombart, T. & Collins, C. (2017) A tutorial for Discriminant Analysis of Principal Components (DAPC) using adegenet 2.1.0. Available from: https://adegenet.r-forge.r-project.org/files/tutorial-dapc.pdf (accessed 10 October 2023)
  52. Kiew, B.H. (1984) Rana malesiana, a new frog from the Sunda Region. Malayan Nature Journal, 37, 151–161.
  53. Kiew, B.H. (1987) An annotated checklist of the herpetofauna of Ulu Endau, Johor, Malaysia. Malayan Nature Journal, 41 (2–3), 413–424.
  54. Köhler, G., Zwitzers, B., Than, N.L., Gupta, D.K., Janke, A., Pauls, S.U. & Thammachoti, P. (2021) Bioacoustics reveal hidden diversity in frogs: Two new species of the genus Limnonectes from Myanmar (Amphibia, Anura, Dicroglossidae). Diversity, 13 (9), 399. https://doi.org/10.3390/d13090399
  55. Lanfear, R., Frandsen, P.B., Wright, A.M., Senfeld, T. & Calcott, B. (2017) PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses. Molecular Biology and Evolution, 34 (3), 772–773. https://doi.org/10.1093/molbev/msw260
  56. Leong, T.M. (2005) Larval systematics of the Peninsular Malaysian Ranidae (Amphibia: Anura). PhD Thesis, Biological Sciences, National University of Singapore, Singapore, 203 pp.
  57. Leong, T.M., Chan-Ard, T. & Chuaynkern, Y. (2003) Additional Anuran and Saurian Records for Phuket, South Thailand. Tropical Natural History, 3 (1), 17–21. https://doi.org/10.58837/tnh.3.1.102847
  58. Lim, K.K.P. (1989) The amphibian fauna of the Ulu Kinchin, Pahang, Malaysia. Malayan Nature Journal, 43, 322–325.
  59. Manthey, U. & Grossmann, W. (1997) Amphibien & Reptilien Südostasiens. Natur-und Tier-Verlag, Münster, 512 pp.
  60. Matsui, M. & Nishikawa, K. (2014) Description of a new species of Limnonectes from Sarawak, Malaysian Borneo (Dicroglossidae, Anura). Current Herpetology, 33 (2), 135–147. https://doi.org/10.5358/hsj.33.135.
  61. Matsui, M., Belabut, D.M. & Ahmad, N. (2014) Two new species of fanged frogs from Peninsular Malaysia (Anura: Dicroglossidae). Zootaxa, 3881 (1), 75–93. https://doi.org/10.11646/zootaxa.3881.1.6
  62. Matsui, M., Eto, K., Belabut, D.M. & Nishikawa, K. (2017) A New Kalophrynus (Amphibia, Anura, Microhylidae) from Peninsular Malaysia. Current Herpetology, 36 (2), 75–86. https://doi.org/10.5358/hsj.36.75
  63. Matsui, M., Kuraishi, N., Eto, K., Hamidy, A., Nishikawa, K., Shimada, T., Yambun, P., Vairappan, C.S. & Hossman, M.B. (2016) Unusually high genetic diversity in the Bornean Limnonectes kuhlii-like fanged frogs (Anura: Dicroglossidae). Molecular Phylogenetics and Evolution, 102, 305–319. https://doi.org/10.1016/j.ympev.2016.06.009
  64. Matsui, M. (2019) A New Species of Pelophryne from Malay Peninsula (Anura, Bufonidae). Current Herpetology, 30 (2), 128–139. https://doi.org/10.5358/hsj.38.128
  65. Minh, B.Q., Nguyen, M.T. & von Haeseler, A. (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30, 1188–1195. https://doi.org/10.1093/molbev/mst024
  66. Mohd Izam, N.A., Ahmad, N., Md Nor, S., Nazron, N.S., Muhammad Nazir, N.A., Magintan, D., Farinordin, F.A., Amat Darbis, N.D. & Zainul Abidin, M.K. (2021) Herpetofauna diversity in Sg. Deka, Hulu Terengganu, Peninsular Malaysia. Gading Journal of Science and Technology, 4 (1), 86–93. [https://ir.uitm.edu.my/id/eprint/46257/1/46257.pdf]
  67. Mulcahy, D.G., Lee, J.L., Miller, A.H., Chand, M., Thura, M.K. & Zug, G.R. (2018) Filling the BINs of life: Report of an amphibian and reptile survey of the Tanintharyi (Tenasserim) Region of Myanmar, with DNA barcode data. ZooKeys, 757, 85–152. https://doi.org/10.3897/zookeys.757.24453
  68. Nidup, T., Wangkulangkul, S., Satasook, C., Bates, P. & Juthong, W. (2013) Diversity of frog (Amphibia: Anura) and the correlation of habitat heterogeneity in Tarutao Island, Thailand. Proceeding of World Biodiversity Congress, Chiang Mai, Thailand, 2013, 1–8.
  69. Norhayati, A. (2017) Frogs and Toads of Malaysia. Penerbit UKM, Bangi, 101 pp.
  70. Nur Amalina, M.I., Azhari, M., Norshaqinah, A., Nor Azrin, N.A., Shukor, M.N., Aisah, M.S., Amirrudin, A., Grismer, L.L. & Norhayati, A. (2017) Species composition of amphibians and reptiles in Tembat Forest Reserve, Hulu Terengganu, Terengganu, Peninsular Malaysia. Malaysian Applied Biology, 46 (4), 119–129.
  71. Ohler, A., Grosjean, S. & Hoyos, J.M. (1999) Observation de la construction d’un nid chez Taylorana hascheana (Anura: Ranidae). Revue française d’aquariologie, Nancy, 26 (1–2), 67–70.
  72. Oksanen, J., Blanchet, F.G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P.R., O’Hara, R.B., Simpson, G.L., Solymos, P., Stevens, M.H.H., Szoecs, E. & Wagner, H. (2020) vegan: Community Ecology Package. R package version 2.5-7. Available from: https://cran.r-project.org/web/packages/vegan/ (accessed 20 January 2026)
  73. Ospina-Sarria, J.J. & Duellman, W.E. (2019) Two new species of Pristimantis (Amphibia: Anura: Strabomantidae) from southwestern Colombia. Herpetologica, 75 (1), 85–95. https://doi.org/10.1655/D-18-00019
  74. Palumbi, S.R. (1996) Nucleic acids II: The polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular systematics. Sinauer, Sunderland, Massachusetts, pp. 205–247.
  75. Pauwels, O.S., Laohawat, O.A., Naaktae, W., Puangjit, C., Wisutharom, T., Chimsunchart, C. & David, P. (2002) Reptile and amphibian diversity in Phang-Nga Province, Southern Thailand. Tropical Natural History, 2 (1), 25–30. https://doi.org/10.58837/tnh.2.1.102862
  76. Poyarkov, N.A., Van Nguyen, T., Popov, E.S., Geissler, P., Pawangkhanant, P., Neang, T., Suwannapoom, C. & Orlov, N.L. (2021) Recent progress in taxonomic studies, biogeographic analysis, and revised checklist of amphibians in Indochina. Russian Journal of Herpetology, 28 (3A), 1–100. https://doi.org/10.30906/1026-2296-2021-28-3A-1-110
  77. Quah, E.S.H., Anuar, S., Muin, M.A., Rahman, N., Mustafa, F.S. & Grismer, L.L. (2013) Species diversity of Herpetofauna of Bukit Panchor State Park, Penang, Peninsular Malaysia. Malayan Nature Journal, 64 (4), 193–211.
  78. Quah, E.S., Anuar, S., Grismer, L., Wood, P.L.J., Azizah, S.M. & Muin, M.A. (2017) A new species of frog of the genus Abavorana Oliver, Prendini, Kraus & Raxworthy 2015 (Anura: Ranidae) from Gunung Jerai, Kedah, northwestern Peninsular Malaysia. Zootaxa, 4320 (2), 272–288. https://doi.org/10.11646/zootaxa.4320.2.4
  79. Quah, E.S.H., Wood, P.L.J., Anuar, S. & Muin, M.A. (2020) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia with an updated checklist of the herpetofauna of Tuba Island. Zootaxa, 4767 (1), 138–160. https://doi.org/10.11646/zootaxa.4767.1.6
  80. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67 (5), 901–904. https://doi.org/10.1093/sysbio/syy032
  81. Reilly, S.B., Stubbs, A.L., Karin, B.R., Bi, K., Arida, E., Iskandar, D.T. & McGuire, J.A. (2019) Leap‐frog dispersal and mitochondrial introgression: Phylogenomics and biogeography of Limnonectes fanged frogs in the Lesser Sundas Archipelago of Wallacea. Journal of Biogeography, 46 (4), 757–769. https://doi.org/10.1111/jbi.13526
  82. Ron, S.R., Carrión, J., Caminer, M.A., Sagredo, Y., Navarrete, M.J., Ortega, J.A., Varela-Jaramillo, A., Maldonado-Castro, G.A. & Terán, C. (2020) Three new species of frogs of the genus Pristimantis (Anura, Strabomantidae) with a redefinition of the P. lacrimosus species group. ZooKeys, 993, 121–155. https://doi.org/10.3897/zookeys.993.53559
  83. Ruzman, N.H.A., Azman, M.S., Fauzi, N.F.M., Munian, K., Mahyudin, N.A.A., Azahar, M.A., Baharudin, M.S., Appanan, M., Beri, A.Z.Z. & Tahir, R. (2023) Small vertebrates in CFS ecological corridor (D-PL1: Lojing–Sungai Brok-Sungai Betis Forest Reserves) Kelantan. BIO Web of Conferences, 73 (01010), 9. https://doi.org/10.1051/bioconf/20237301010
  84. Savage, J.M. & Heyer, W.R. (1997) Digital webbing formulae for anurans: a refinement. Herpetological Review, 28 (3), 131.
  85. Sclater, W.L. (1892) On some specimens of frogs in the Indian Museum, Calcutta, with descriptions of several new species. Proceedings of the Zoological Society of London, 1892, 341–348.
  86. Sheridan, J.A. & Stuart, B.L. (2018) Hidden species diversity in Sylvirana nigrovittata (Amphibia: Ranidae) highlights the importance of taxonomic revisions in biodiversity conservation. PLoS One, 13 (4), e0196242. https://doi.org/10.1371/journal.pone.0192766
  87. Stuart, B.L., Inger, R.F. & Voris, H.K. (2006) High level of cryptic species diversity revealed by sympatric lineages of Southeast Asian Forest frogs. Biology Letters, 2 (3), 470–474. https://doi.org/10.1098/rsbl.2006.0505
  88. Sukumaran, J. (2002) The amphibian fauna of a forested area in Temengor, Perak, Malaysia, with the first record of Philautus parvulus (Boulenger, 1893) in the Malay Peninsula. Hamadryad, 27, 1–10.
  89. Sumarli, A.X., Grismer, L.L., Anuar, S., Muin, M.A. & Quah, E.S.H. (2015) First report on the amphibians and reptiles of a remote mountain, Gunung Tebu in northeastern Peninsular Malaysia. Check List, 11 (4), 1–32. https://doi.org/10.15560/11.4.1679
  90. Syafiq, M.F., Badli-Sham, B.H., Aqmal-Naser, M., Fahmi-Ahmad, M., Bin Syed Abd Ghani, S.A.F., Mohd A Wahid, M.I., Amir, F.H, Kaliyappan, G., Sapri, N.N., Nordin, N.F., Azzimin, J., Shahril-Ridhwan, M., Kei, L.T., Shu, X.N., Rizal, S.A & Ahmad, A. (2024) Herpetofauna diversity of the disturbed and isolated Bukit Maras in Terengganu, Peninsular Malaysia. ARPHA Preprints. Available from: https://preprints.arphahub.com/article/122806/ (accessed 20 January 2026) https://doi.org/10.3897/arphapreprints.e122806
  91. Tamura, K., Stecher, G. & Kumar, S. (2021) MEGA11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology And Evolution, 38 (7), 3022–3027. https://doi.org/10.1093/molbev/msab120
  92. Thong-aree, S., Chan-ard, T., Cota, M. & Makchai, S. (2011) The amphibian fauna of Bala Forest reported from southernmost Thailand in 2011. Thailand Natural History Museum Journal, 5 (2), 99–106.
  93. Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44 (W1), 232–235. https://doi.org/10.1093/nar/gkw256
  94. Trofimets, A.V., Dufresnes, C., Pawangkhanant, P., Bragin, A.M., Gorin, V.A., Hasan, M., Lalremsanga, H.T., Muin, M.A., Le, D.X., Van Nguyen, T. & Suwannapoom, C. (2024) Four in one: An integrative taxonomic revision of the Microhyla berdmorei complex (Amphibia: Anura: Microhylidae) illustrates the tremendous amphibian diversity of Southeast Asia. Vertebrate Zoology, 74, 595–641. https://doi.org/10.3897/vz.74.e127937
  95. Van Kampen, P.N. (1923) The amphibia of the Indo-Australian archipelago. E. J. Brill Ltd., Leiden, 304 pp. https://doi.org/10.1163/9789004590182
  96. Visoot, Y., Makchai, S., Khonsue, W., Traijitt, T., Srion, L., Kitana, N., Kitana, J. & Thammachoti, P. (2023) Herpetofauna of Yao Islands, Phang-nga Province, Southern Thailand: A list of species and habitat notes. Thailand Natural History Museum Journal, 17, 29–43.
  97. Wilcox, T.P., Zwickl, D.J., Heath, T.A. & Hillis, D.M. (2002) Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution, 25 (2), 361–371.
  98. Wood, P.L.J., Grismer, L.L., Youmans, T.M., Nasir, N.B., Ahmad, N. & Senwai, J. (2008) Additions to the herpetofauna of Endau-Rompin, Johor, west Malaysia. Herpetological Review, 39 (1), 112–121.
  99. Yari, P., Valiee, T., Bashiri, H., Kahrizi, D. & Yari, K. (2017) An efficient and optimized protocol for DNA extraction from animal tissues. Biological, Environmental and Agricultural Sciences, 2, 41–44.
  100. Yodthong, S., Rujirawan, A., Stuart, B.L. & Aowphol, A. (2021) A new Limnonectes (Anura: Dicroglossidae) from Southern Thailand. Animals, 11 (2), 566. https://doi.org/10.3390/ani11020566
  101. Yodthong, S., Stuart, B.L. & 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
  102. Zhang, P., Liang, D., Mao, R.L., Hillis, D.M., Wake, D.B. & Cannatella, D.C. (2013) Efficient sequencing of anuran mtDNAs and a mitogenomic exploration of the phylogeny and evolution of frogs. Molecular Biology and Evolution, 30 (8), 1899–1915. https://doi.org/10.1093/molbev/mst091
  103. Ziegler, T. (2002) Die Amphibien und Reptilien eines Tieflandfeuchtwald-Schutzgebietes in Vietnam. Natur and Tier-Verlag, Münster, 342 pp.
  104. Ziegler, T. & Nguyen, T.Q. (2021) New records and an updated checklist of amphibians and snakes from Tuyen Quang Province, Vietnam. Bonn Zoological Bulletin, 70 (1), 201–219. https://doi.org/10.20363/BZB-2021.70.1.201
  105. Zou, B., Anuar, S., Low, T.J., Hong, Z., Grismer, L.L. & Quah, E.S.H. (2024) Advertisement calls from species of the Limnonectes hascheanus-limborgi complex (Anura: Dicroglossidae) from Peninsular Malaysia. Zootaxa, 5399 (2), 163–171. https://doi.org/10.11646/zootaxa.5399.2.5

How to Cite

Zou, B., Anuar, M.S., Low, T.J., Hong, Z., Chau, E., Grismer, L.L. & Quah, E.S.H. (2026) A systematic review of the Limnonectes hascheanus-limborgi complex (Anura: Dicroglossidae), with the description of a new species from Peninsular Malaysia. Zootaxa, 5785 (1), 67–102. https://doi.org/10.11646/zootaxa.5785.1.3