Abstract
A new species of Amolops is described from Vu Quang National Park, Ha Tinh Province, North-central Vietnam based on morphological and molecular differences. Morphologically, Amolops minhlei sp. nov. is distinguishable from its congeners by a combination of the following morphological characteristics: SVL 31.8–35.8 mm in males, 36.7–45.8 mm in females; head longer than wide; vomerine teeth absent; snout long (SE/SVL 0.16–0.17 in males, 0.16 in females); tympanum distinct, round (TD/ED 0.41–0.42 in males, 0.40–0.43 in females); skin smooth; supratympanic fold indistinct; dorsolateral fold present; webbing formula I0–0II0–1/2III0–1/2IV1/2–0V; dorsum olive green or reddish brown with a few dark spots; a brown dorsolateral stripe extending from behind eye to above vent; dorsal surface of fore- and hind-limbs light grey with brown crossbars, throat, chest and belly white; males with external vocal sac and nuptial pad on finger I; and flanks grey brown with small white glandular spot. In phylogenetic analyses, the new species is strongly supported as a member of the A. monticola group. However, the new species differs from other members of the A. monticola group by 10.00–19.66% pairwise genetic divergence in the mitochondrial ND2 gene fragment and 1.56–6.67% in the mitochondrial 16S gene fragment.
References
- Bain, R.H., Lathrop, A., Murphy, R.W., Orlov, N.L. & Ho, C.T. (2003) Cryptic species of a cascade frog from Southeast Asia: taxonomic revisions and descriptions of six new species. American Museum Novitates, 3417, 1–60. https://doi.org/10.1206/0003-0082(2003)417%3C0001:CSOACF%3E2.0.CO;2
- Bain, R.H. & Nguyen, T.Q. (2004) Herpetofaunal diversity of Ha Giang Province in northeastern Vietnam, with descriptions of two new species. American Museum Novitates, 3453, 1–42. https://doi.org/10.1206/0003-0082(2004)453%3C0001:HDOHGP%3E2.0.CO;2
- Bain, R.H., Stuart, B.L. & Orlov, N.L. (2006) Three new Indochinese species of cascade frogs (Amphibia: Ranidae) allied to Rana archotaphus. Copeia, 2006, 43–59. https://doi.org/10.1643/0045-8511(2006)006[0043:TNISOC]2.0.CO;2
- Biju, S., Mahony, S. & Kamei, R.G. (2010) Description of two new species of torrent frog, Amolops Cope (Anura: Ranidae) from a degrading forest in the northeast Indian state of Nagaland. Zootaxa, 2408 (1), 31–66. https://doi.org/10.11646/zootaxa.2408.1.2
- Boulenger, G.A. (1903) Descriptions of three new batrachians from Tonkin. Annals and Magazine of Natural History, Series 7, 12, 186–188. https://doi.org/10.1080/00222930308678835
- Boulenger, G.A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum, 20, 1–226. https://doi.org/10.26515/rzsi/v20/i1/1920/163533
- 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
- Che, J., Jiang, K., Yan, F. & Zhang, Y. (2020) Amphibians and Reptiles in Tibet Diversity and Evolution. Chinese Academy of Sciences, Science Press, Beijing, 803 pp. [in Chinese with English abstracts and species descriptions]
- Cope, E.D. (1865) Sketch of the primary groups of Batrachia s. Salientia. Natural History Review, New Series 5, 97–120.
- Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jmodelTest 2: More models, new heuristics and high–performance computing. Nature Methods, 9, e772. https://doi.org/10.1038/nmeth.2109
- Fei, L., Hu, S.Q., Ye, C.Y. & Huang, Y.Z. (2009) Fauna Sinica. Amphibia. Vol. 3 Anura. Science Press, Beijing, 881 pp.
- Fei, L., Ye, C.Y. & Jiang, J.P. (2010) Colored atlas of Chinese amphibians. Sichuan Publishing Group, Sichuan Publishing House of Science & Technology, Sichuan, 519 pp.
- Frost, D.R. (2025) Amphibian species of the world version 6.2, an online reference. Available from: http://research.amnh.org/vz/herpetology/amphibia/ (accessed 12 August 2025)
- Gan, Y.-L., Qin, T., Lwin, Y.-H., Li, G.-G., Quan, R.-C., Liu, S. & Yu, G.-H. (2020a) A new species of Amolops (Anura: Ranidae) from northern Myanmar. Zoological Research/Dōngwùxué yánjiū. Kunming, 41, 733−739. https://doi.org/10.24272/j.issn.2095-8137.2020.018
- Gan, Y.-L., Yu, G. & Wu, Z. (2020b) A new species of the genus Amolops (Anura: Ranidae) from Yunnan, China. Zoological Research, 41 (2), 188−193. https://doi.org/10.24272/j.issn.2095-8137.2020.125
- Glaw, F. & Vences, M. (2007) A field guide to the amphibians and reptiles of Madagascar. 3rd Edition. Frosch Verlag, Cologne, 495 pp.
- Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematics Biology, 42 (2), 182–192. https://doi.org/10.1093/sysbio/42.2.182
- Hoang, C.V., Jiang, J.-P., Nguyen, H.Q., Orlov, N.L., Le, M.V., Nguyen, S.H.L., Nguyen, V.H., Nguyen, T.T. & Ziegler, T. (2021) A new species of Vietnamophryne from Vietnam. Revue Suisse de Zoologie, 128, 207–219. https://doi.org/10.35929/RSZ.0046
- Hoang, D.T., Chernomor, O., Von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: Improving the Ultrafast Bootstrap Approximation. Molecular Biology and Evolution, 35 (2), 518–522. https://doi.org/10.1093/molbev/msx281
- Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42, 182–192. https://doi.org/10.1093/sysbio/42.2.182
- Inger, R.F. & Chanard, T. (1997) A new species of ranid frog from Thailand, with comments on Rana livida (Blyth). Natural History Bulletin of the Siam Society, 45, 65–70.
- IUCN (2025) The IUCN Red List of Threatened Species. Version 2025-2. Available from: https://www.iucnredlist.org (accessed 28 December 2025)
- Jiang, K., Fang, Y., Jiang, X., Zou, D.H., Jiang, J.P. & Che, J. (2016) A new species of the genus Amolops (Amphibia: Ranidae) from southeastern Tibet, China. Zoological Research, 37, 31–40.
- Jiang, K., Ren, J.-L., Lyu, Z.-T., Wang, D.Z., Wang, K.-L., Wu, J.-W. & Li, J.-T. (2021) Taxonomic revision of Amolops chunganensis (Pope, 1929) (Amphibia: Anura) and description of a new species from southwestern China, with discussion on Amolops monticola group and assignment of species groups of the genus Amolops. Zoological Research. Kunming, 42, 574–591. https://doi.org/10.24272/j.issn.2095-8137.2021.107
- Jombart, T., Devillard, S. & Balloux, F. (2010) Discriminant analysis of principal components: a new method for the analysis of genetically structured populations. BMC Genetics, 11, 1–15. [http://www.biomedcentral.com/1471-2156/11/94] https://doi.org/10.1186/1471-2156-11-94
- Lanfear, R.B., Calcott, S.Y.H. & Guindon, S. (2012) Partition finder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology Evolution, 29, 1695–701. https://doi.org/10.1093/molbev/mss020
- Liu, W.Z., Yang, D.T., Ferraris, C., Matsui, M. & Price, A.H. (2000) Amolops bellulus: A new species of stream-breeding frog from western Yunnan, China (Anura: Ranidae). Copeia, 2000, 536–541. https://doi.org/10.1643/0045-8511(2000)000[0536:ABANSO]2.0.CO;2
- Liu, S., Hou, M., Mo, M., Lu, Y., Guo, J., Wang, W., Zhang, W. & Rao, D. (2025) A new species of the Amolops monticola group (Anura, Ranidae) from southern Yunnan, China. Journal of Asia-Pacific Biodiversity, 18, 258–268. https://doi.org/10.1016/j.japb.2024.08.002
- Lleonart, J., Salat, J. & Torres, G.J. (2000) Removing allometric effects of body size in morphological analysis. Journal of Theoretical Biology, 205, 85–93. https://doi.org/10.1006/jtbi.2000.2043
- Nguyen, L.T., Tapley, B., La, V.T. & Rowley, J.J.L. (2025) A new species of Amolops monticola group (Amphibia: Ranidae) from Hoang Lien Range, northwest Vietnam. Zootaxa, 5594 (3), 485–508. https://doi.org/10.11646/zootaxa.5594.3.3
- Ohler, A., Wollenberg, K.C., Grosjean, S., Hendrix, R., Vences, M., Ziegler, T. & Dubois, A. (2011) Sorting out Lalos: description of new species and additional taxonomic data on megophryid frogs from northern Indochina (genus Leptolalax, Megophryidae, Anura). Zootaxa, 3147 (1), 1–83. https://doi.org/10.11646/zootaxa.3147.1.1
- Orlov, N.L., Ananjeva, N.B., Ermakov, O.A., Lukonina, S.A., Ninh, T.H. & Nguyen, T.T. (2024) A new record of Bufo gargarizans Complex (Bufonidae, Anura) from Truong Son Mounts, Ha Tinh and Ha Giang Provinces, Vietnam based on molecular evidence with a description of a new species. Diversity, 16 (361), 1–16. https://doi.org/10.3390/d16070361
- Patel, N.G., Garg, S., Das, A., Stuart, B.L. & Biju, S.D. (2021) Phylogenetic position of the poorly known montane cascade frog Amolops monticola (Ranidae) and description of a new closely related species from Northeast India. Journal of Natural History, 55, 1403–1440. https://doi.org/10.1080/00222933.2021.1946185
- Pham, C.T., Le, M.D., Hoang, C.V., Pham, V.A., Ziegler, T. & Nguyen, Q.T. (2020) First records of Bufo luchunnicus (Yang et Rao, 2008) and Amolops wenshanensis Yuan, Jin, Li, Stuart et Wu, 2018 (Anura: Bufonidae, Ranidae) from Vietnam. Russian Journal of Herpetology, 27, 81–86. https://doi.org/10.30906/1026-2296-2021-28-4-219-224
- Pham, A.V., Hoang, C.V., Tapley, B., Nguyen, T.L., Nguyen, H.H., La, V.T., Ziegler, T., Rowley, J.J.L., Nguyen, Q.T. & Le, D.M. (2025) Amolops cuongi (Amphibia, Anura, Ranidae), a new species from the Hoang Lien Range, Vietnam. ZooKeys, 1256, 235–257. https://doi.org/10.3897/zookeys.1256.158846
- Pham, A.V., Pham, C.T., Ngo, H.T., Sung, N.B., Ziegler, T. & Le, M.D. (2023) A new species of Amolops (Anura: Ranidae) from Son La Province, northwestern Vietnam. Raffles Bulletin of Zoology, 71, 59–69. https://doi.org/10.26107/RBZ-2023-0004
- Pope, C.H. (1929) Four new frogs from Fukien Province, China. American Museum Novitates, 352, 1–5.
- Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
- R Core Team (2025) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: http://www.R–project.org (accessed 25 November 2025)
- Ray, P. (1992) Two new hill-stream frogs of the genus Amolops Cope (Amphibia: Ranidae) from Uttar Pradesh (India). Indian Journal of Forestry, 15, 345–350.
- Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61 (3), 539−542. https://doi.org/10.1093/sysbio/sys029
- Simmons, J.E. (2002) Herpetological collecting and collections management. Revised edition. Society for the Study of Amphibians and Reptiles, Herpetological Circular, 31, 1–153.
- Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomologial Society America, 87 (6), 651–701. https://doi.org/10.1093/aesa/87.6.651
- Smith, M.A. (1940) The amphibians and reptiles obtained by Mr. Ronald Kaulback in Upper Burma. Records of the Indian Museum, 42, 465–486. https://doi.org/10.26515/rzsi/v42/i3/1940/162431
- 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, 470–474. https://doi.org/10.1098/rsbl.2006.0505
- Stuart, B.L., Bain, R.H., Phimmachak, S. & Spence, K. (2010) Phylogenetic systematics of the Amolops monticola group (Amphibia: Ranidae), with description of a new species from northwestern Laos. Herpetologica, 66, 52–66. https://doi.org/10.1655/08-073.1
- Swofford, D.L. (2001) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. [program]
- Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The ClustalX windows interface: Xexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Research, 25, 4876−4882. https://doi.org/10.1093/nar/25.24.4876
- Thorpe, R.S. (1975) Quantitative handling of characters useful in snake systematics with particular reference to interspecific variation in the Ringed Snake Natrix natrix (L.). Biological Journal of the Linnean Society, 7, 27–43. https://doi.org/10.1111/j.1095-8312.1975.tb00732.x
- Thorpe, R.S. (1983) A review of the numerical methods for recognizing and analyzing racial differentiation. In: Felsenstein, J. (Ed.), Numerical Taxonomy. NATO ASI Series (Series G: Ecological Sciences). Vol. 1. Springer, Berlin, pp. 404–423. https://doi.org/10.1007/978-3-642-69024-2_43
- Tordoff, A.W., Tran, B.Q., Nguyen, T.D. & Le, H.M. (Eds.) (2004) Sourcebook of existing and proposed protected areas in Vietnam. 2nd Edition. Birdlife International in Indochina and Ministry of Agriculture and Rural Development, Hanoi. [CD-Rom]
- Turan, C. (1999) A note on the examination of morphometric differentiation among fish populations: The Truss System. Turkish Journal of Zoology, 23, 259–263.
- Wang, J., Li, J., Du, L., Hou, M. & Yu, G. (2022) A cryptic species of the Amolops ricketti species group (Anura, Ranidae) from China–Vietnam border regions. ZooKeys, 1112, 139–159. https://doi.org/10.3897/zookeys.1112.82551
- Wong, T.K.F., Ly-Trong, N., Ren, H., Banos, H., Roger, A.J., Susko, E., Bielow, C., De Maio, N., Goldman, N., Hahn, M.W., Huttley, G., Lanfear, R. & Minh, B.Q. (2025) IQ-TREE 3: Phylogenomic Inference Software using Complex Evolutionary Models. Submitted. Available from: https://ecoevorxiv.org/repository/view/8916/ (accessed 12 January 2026) https://doi.org/10.32942/X2P62N
- Wu, G.F. & Tian, W.S. (1995) A new Amolops species from southern Yunnan. Sichuan Journal of Zoology (Supplement), Herpetological Series, 8, 50–52.
- Ye, C.Y., Fei, L. & Hu, S.Q. (1993) Rare and Economic Amphibians of China. Sichuan Publishing House of Science and Technology, Chengdu, 412 pp.
- Yu, G., Wu, Z. & Yang, J.X. (2019) A new species of the Amolops monticola group (Anura: Ranidae) from southwestern Yunnan, China. Zootaxa, 4577 (3), 548–560. https://doi.org/10.11646/zootaxa.4577.3.8
- Yuan, Z.Y., Jin, J.Q., Li, J.N., Stuart, B.L. & Wu, J. (2018) A new species of cascade frog (Amphibia: Ranidae) in the Amolops monticola group from China. Zootaxa, 4415 (3), 498–512. https://doi.org/10.11646/zootaxa.4415.3.5
- Zhao, W.G., Rao, D.Q., Lü, S.Q. & Dong, B.J. (2005) Herpetological surveys of Xizang Autonomous Region 2. Mêdog. Sichuan Journal of Zoology, 24, 250–253.
