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Type: Article
Published: 2023-11-10
Page range: 301-335
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Two new species of Leptobrachella Smith 1925 (Amphibia: Megophryidae) from Cao Bang Province, Vietnam

Institute of Ecology and Biological Resources; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Hanoi 10072; Vietnam
Institute of Ecology and Biological Resources; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Hanoi 10072; Vietnam; Forest resources and environment center; 300 Ngoc Hoi Road; Thanh Tri; Hanoi; Vietnam
Institute of Ecology and Biological Resources; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Hanoi 10072; Vietnam; Graduate University of Science and Technology; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Cau Giay; Hanoi 10072; Vietnam
Zoologischer Garten Köln; Riehler Str. 173; D-50735 Cologne; Germany; Institute of Zoology; University of Cologne; Zülpicher Str. 47b; D–50674 Cologne; Germany
Institute of Ecology and Biological Resources; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Hanoi 10072; Vietnam; Graduate University of Science and Technology; Vietnam Academy of Science and Technology; 18 Hoang Quoc Viet Road; Cau Giay; Hanoi 10072; Vietnam
Amphibia Leptobrachella phiaoacensis sp. nov. Leptobrachella phiadenensis sp. nov. Phia Oac-Phia Den National Park phylogenetics taxonomy

Abstract

Two new species, Leptobrachella phiaoacensis sp. nov. and Leptobrachella phiadenensis sp. nov., are described from Phia Oac-Phia Den National Park, Cao Bang Province, Vietnam based on genetic divergence and morphological differences. Leptobrachella phiaoacensis sp. nov. is characterized by a combination of the following characters: size medium (27.8–33.3 mm in five adult males and 31.5–41.8 mm in five adult females); head longer than wide; tympanum distinct; dorsal skin shagreened with fine tubercles; toes webbing rudimentary, with narrow lateral fringes; supratympanic fold edged by a distinct black line; the presence of a dark brown triangle in interorbital region and a W-shaped marking in scapular region; belly from pinkish white to white with dark brown specking on belly periphery; iris bicolored copper in upper part, fading to silvery grey in lower part. Leptobrachella phiadenensis sp. nov. is distinguished from its congeners by a combination of the following morphological characters: size medium (27.6–28.6 mm in two adult females); head wider than long; tympanum distinct; skin on dorsal head and body rough with warts, tubercles and dermal ridges; toes webbing rudimentary, with narrow dermal fringes; supratympanic fold orange; dorsal surface of head and body with some large dark brown markings, a dark brown triangle in interorbital region, edged in white; flank light brown with some dark flecks; belly white with dark specking on outer margins; iris bicolored, copper in upper part, fading to silvery grey in lower part. In terms of genetic distance, the two new species differ from each other and other congeners for which comparable sequences are available by at least 3.7% (16S rRNA gene). Recognition of these new species brings the total number of known species in the genus Leptobrachella from Vietnam to 34.

 

References

  1. Anderson, J. (1871) A list of the reptilian accession to the Indian Museum, Calcutta, from 1865 to 1870, with a description of some new species. Journal of the Asiatic Society of Bengal, 40, 12–39.
  2. Boulenger, G.A. (1893) Concluding report on the reptiles and batrachians obtained in Burma by Signor L. Fea, dealing with the collection made in Pegu and the Karin Hills in 1887–88. Annali del Museuo Civico di Genoa, 13, 304–347.
  3. Chen, J.M., Poyarkov, N.A. Jr., Suwannapoom, C., Lathrop, A., Wu, Y.H., Zhou, W.W., Yuan, Z.Y., Jin, J.Q., Chen, H.M., Liu, H.Q., Nguyen, T.Q., Nguyen, S.N., Duong, T.V., Eto, K., Nishikawa, K., Matsui, M., Orlov, N.L., Stuart, B.L., Brown, R.M., Rowley, J.J.L., Murphy, R.W., Wang, Y.Y. & Che, J. (2018) Large-scale phylogenetic analyses provide insights into unrecognized diversity and historical biogeography of Asian leaf-litter frogs, genus Leptolalax (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 124, 162–171. https://doi.org/10.1016/j.ympev.2018.02.020
  4. Chen, J.M., Xu, K., Poyarkov, N.A., Wang, Jr.K., Yuan, Z.Y., Hou, M., Suwannapoom, C., Wang, J. & Che, J. (2020) How little is known about “the little brown frogs”: description of three new species of the genus Leptobrachella (Anura: Megophryidae) from Yunnan Province, China. Zoological Research, 41, 1–22. https://doi.org/10.24272/j.issn.2095-8137.2020.036
  5. Chen, W.C., Liao, X.W., Zhou, S.C. & Mo, Y.M. (2019) A new species of Leptobrachella (Anura: Megophryidae) from southern Guangxi, China. Zootaxa, 4563 (1), 67–82. https://doi.org/10.11646/zootaxa.4563.1.3
  6. Chen, W.C., Yu, G.D., Cheng, Z.Y., Meng, T., Wei, H., Zhou, G.Y. & Lu, Y.W. (2021) A new species of Leptobrachella (Anura: Megophryidae) from central Guangxi, China. Zoological Research, 42, 783–788. https://doi.org/10.24272/j.issn.2095-8137.2021.179
  7. Cheng, Y.L., Shi, S., Li, J., Liu, J., Li, S.Z. & Wang, B. (2021) A new species of the Asian leaf litter toad genus Leptobrachella Smith, 1925 (Anura, Megophryidae) from northwest Guizhou Province, China. ZooKeys, 1021, 81–107. https://doi.org/10.3897/zookeys.1021.60729
  8. Das, I., Lyngdoh, T.R.K., Rangad, D. & Hooroo, R.N.K. (2010) A new species of Leptolalax (Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya, northeastern India. Zootaxa, 2339 (1), 44–56. https://doi.org/10.11646/zootaxa.2339.1.2
  9. Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24, 6–21.
  10. Dubois, A. (1983) Note préliminaire sur le genre Leptolalax Dubois, 1980 (Amphibiens, Anoures), avec diagnose d'une espèce nouvelle du Vietnam. Alytes, 2, 147–153. https://doi.org/10.5962/p.297429
  11. Duong, T.V., Do, D.T., Ngo, C.D., Nguyen, T.Q. & Poyarkov, N.A. (2018) A new species of the genus Leptolalax (Anura: Megophryidae) from southern Vietnam. Zoological Research, 39, 181–196. https://doi.org/10.24272/j.issn.2095-8137.2018.009
  12. Eto, K., Matsui, M. & Nishikawa, K. (2015) Description of a new species of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from Borneo. Current Herpetology, 34, 128–139. https://doi.org/10.5358/hsj.34.128
  13. Eto, K., Matsui, M. & Nishikawa, K. (2016) A new highland species of dwarf litter frog genus Leptobrachella (Amphibia: Anura: Megophryidae) from Sarawak. Raffles Bulletin of Zoology, 64, 194–203
  14. Fei, L., Hu, S.Q., Ye, C.Y. & Huang, Y.Z. (2009) Fauna Sinica. Amphibia. Vol. 2. Anura. Science Press, Beijing, 957 pp.
  15. Fei, L., Ye, C. & Huang, Y. (1991 “1990”) Key to Chinese Amphibia. Chongqing Branch Science and Technology Literature Press, Chongquing, 364 pp.
  16. Fei, L., Ye, C. & Jiang, J. (2012) Colored Atlas of Chinese Amphibians. Sichuan Publishing Group. Sichuan Publishing House of Science & Technology, Sichuan, 620 pp.
  17. Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & & Gemmellet, N.J. (2007) Underestimation of Species Richness in Neotropical Frogs Revealed by mtDNA Analyses. PLoS ONE, 2 (10), e1109. https://doi.org/10.1371/journal.pone.0001109
  18. Frost, D.R. (2023) Amphibian Species of the World 6.0, an Online Reference. Electronic Database accessible. Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 1 February 2023)
  19. Frost, D.R., Grant, T., Faivovich, J., Bain, R., Haas, A., Haddad, C.F.B., de Sa, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–291. https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
  20. Hall, T.A. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
  21. Hoang, C.V., Nguyen, T.T., Luu, V.Q., Nguyen, T.Q. & Jiang, J.P. (2019) A new species of Leptobrachella Smith 1925 (Anura: Megophryidae) from Thanh Hoa Province, Vietnam. Raffles Bulletin of Zoology, 67, 536–556. https://doi.org/10.26107/RBZ-2019-0042.
  22. Hou, Y., Zhang, M., Hu, F., Li, S., Shi, S., Chen, J., Mo, X.Y. & Wang, B. (2018) A new species of the genus Leptolalax (Anura, Megophryidae) from Hunan, China. Zootaxa, 4444 (3), 247–266. https://doi.org/10.11646/zootaxa.4444.3.2
  23. 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
  24. Humtsoe, L.N., Bordoloi, S., Ohler, A. & Dubois, A. (2008) Rediscovery of a long known species, Ixalus lateralis Anderson, 1871. Zootaxa, 1921 (1), 24–34. https://doi.org/10.11646/zootaxa.1921.1.2
  25. Inger, R.F., Orlov, N. & Darevsky, I. (1999) Frogs of Vietnam: a report on new collections. Fieldiana: Zoology, 92, 1–46.
  26. Jiang, K., Yan, F., Suwannapoom, C., Chomdej, S. & Che, J. (2013) A new species of the genus Leptolalax (Anura: Megophryidae) from northern Thailand. Asian Herpetological Research, 4, 100–108. https://doi.org/10.3724/SP.J.1245.2013.00100
  27. Jobb, G. (2011) Treefinder Version of March 2011. Software Distributed by the Author. Available from: http://www.treefinder.de (assessed 10 Febuary 2023)
  28. Lathrop, A., Murphy, R.W., Orlov, N. & Ho, C.T. (1998) Two new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Amphibi-Reptilia, 19, 253–267. https://doi.org/10.1163/156853898X00160
  29. Leaché, A.D. & Reeder, T.W. (2002) Molecular systematics of the eastern fence lizard (Sceloporus undulatus): a comparison of parsimony, likelihood, and Bayesian approaches. Systematic Biology, 51, 44–68. https://doi.org/10.1080/106351502753475871
  30. Li, S.Z., Liu, J., Wei, G. & Wang, B. (2020) A new species of the Asian leaf litter toad genus Leptobrachella (Amphibia, Anura, Megophryidae) from southwest China. ZooKeys, 943, 91–118. https://doi.org/10.3897/zookeys.943.51572
  31. Lin, S.S., Li, Y.H., Lu, Y.H., Su, H.L., Wu, S.B., Zhang, Q.Q., Mo, M.J., Xiao, S.J., Pan, Z., Pan, H.J., Zeng, Z.C. & Wang, J. (2022) A new species of the genus Leptobrachella (Anura, Megophryidae) from northwestern Guangdong Province, China. Herpetozoa, 35, 165–178. https://doi.org/10.3897/herpetozoa.35.e89981
  32. Liu, C. (1950) Amphibians of western China. Fieldiana: Zoology, 2, 1–400. https://doi.org/10.5962/bhl.part.4737
  33. Liu, J., Shi, S., Li, S., Zhang, M., Xiang, S., Wei, G. & Wang, B. (2023) A new Asian leaf litter toad of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from central south China. ZooKeys, 1149, 103–134. https://doi.org/10.3897/zookeys.1149.85895
  34. Lo, N.T., Hoang, C.V., Nguyen, T.Q., Nguyen, S.L.H., Ziegler, T. & Pham, C.T. (2022) First record of Leptebrachella shiwandashanensis Chen, Peng, Pan, Liao, Liu & Huang, 2021 (Anura: Megophryidae) from Vietnam. Journal of Forestry Science and Technology, 14, 28–32. https://doi.org/10.55250/jo.vnuf.2022.14.028-032
  35. Luo, T., Wang, W., Peng, D., Lei, B., Deng, H., Ji, S., Huang, H. & Zhou, J. (2022) A new species of the Asian Leaf Litter Toad genus Leptobrachella (Amphibia, Anura, Megophryidae) from Chongqing City, Southwest China. Asian Herpetological Research, 13, 75–95. https://doi.org/10.16373/j.cnki.ahr.210052
  36. Luo, T., Xiao, N., Gao, K. & Zhou, J. (2020) A new species of Leptobrachella (Anura, Megophryidae) from Guizhou Province, China. ZooKeys, 923, 115–140. https://doi.org/10.3897/zookeys.923.47172.
  37. Lyu, J.C., Dai, L.L., Wei, P.F., He, Y.H., Yuan, Z.Y., Shi, W.L., Zhou, S.L., Ran, S.Y., Kuang, Z.F., Guo, X., Wei, G. & Yuan, G. (2020) A new species of the genus Leptobrachella Smith, 1925 (Anura, Megophryidae) from Guizhou, China. ZooKeys, 1008, 139–157. https://doi.org/10.3897/zookeys.1008.56412
  38. Mahony, S., Foley, N.M., Biju, S.D. & Teeling, E.C. (2017) Evolutionary History of the Asian Horned Frogs (Megophryinae): Integrative Approaches to Timetree Dating in the Absence of a Fossil Record. Molecular Biology and Evolution, 34 (3), 744–771 https://doi.org/10.1093/molbev/msw267
  39. Mathew, R. & Sen, N. (2010 "2009") Description of a new species of Leptobrachium Tschudi, 1838 (Amphibia: Anura: Megophryidae) from Meghalaya, India. Records of the Zoological Survey of India, 109, 91–108.
  40. Matsui, M. (2006) Three new species of Leptolalax from Thailand (Amphibia, Anura, Megophryidae). Zoological Science, 23, 821–830. https://doi.org/10.2108/zsj.23.821
  41. Matsui, M., Eto, K., Nishikawa, K., Hamidy, A., Belabut, D., Ahmad, N., Panha, S., Khonsue, W. & Grismer, L.L. (2017) Mitochondrial Phylogeny of Leptolalax from Malay Peninsula and Leptobrachella (Anura, Megophryidae). Current Herpetology, 36 (1), 11–21. https://doi.org/10.5358/hsj.36.11
  42. Matsui, M., Hamidy, A., Murphy, R.W., Khonsue, W., Yambun, P., Shimada, T., Ahmad, N., Belabut, D.M. & Jiang, J.P. (2010) Phylogenetic relationships of megophryid frogs of the genus Leptobrachium (Amphibia, Anura) as revealed by mtDNA gene sequences. Molecular Phylogenetics and Evolution, 56 (1), 259–272.
  43. Matsui, M., Nishikawa, K. & Yambun, P. (2014) A new Leptolalax from the mountains of Sabah, Borneo (Amphibia, Anura, Megophryidae). Zootaxa, 3753 (5), 440–452. https://doi.org/10.11646/zootaxa.3753.5.3
  44. Nguyen, T.L., Poyarkov, N.A., Le, T.D., Vo, B.D., Ninh, T.H., Duong, V.T., Murphy, R.W. & Nguyen, S.N. (2018) A new species of Leptolalax (Anura: Megophryidae) from Son Tra Peninsula, central Vietnam. Zootaxa, 4388 (10, 1–21. https://doi.org/10.11646/zootaxa.4388.1.1
  45. Nguyen, L.T., Tapley, B., Nguyen, C.T., Luong, H.V. & Rowley, J.J.L. (2021) A new species of Leptobrachella (Anura, Megophryidae) from Mount Pu Ta Leng, northwest Vietnam. Zootaxa, 5016, 301–332. https://doi.org/10.11646/zootaxa.5016.3.1
  46. Oberhummer, E., Barten, C., Schweizer, M., Das, I., Haas, A. & Hertwig, S.T. (2014) Description of the tadpoles of three rare species of megophryid frogs (Amphibia: Anura: Megophryidae) from Gunung Mulu, Sarawak, Malaysia. Zootaxa, 3835 (1), 59–79. https://doi.org/10.11646/zootaxa.3835.1.3
  47. Ohler, A., Marquis, O., Swan, S. & Grosjean, S. (2000) Amphibian biodiversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa, 13, 71–87.
  48. 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
  49. Poyarkov, N.A., Rowley, J.J.L., Gogoleva, S.I., Vassilieva, A.B., Galoyan, E.A. & Orlov, N.L. (2015) A new species of Leptolalax (Anura: Megophryidae) from the western Langbian Plateau, southern Vietnam. Zootaxa, 3931 (2), 221–252. https://doi.org/10.11646/zootaxa.3931.2.3
  50. Qian, T.Y., Xia, X., Cao, Y., Xiao, N.W. & Yang, D.D. (2020) A new species of Leptobrachella (Anura: Megophryidae) Smith, 1925 from Wuling Mountains in Hunan Province, China. Zootaxa, 4816 (4), 491–526. https://doi.org/10.11646/zootaxa.4816.4.4
  51. 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
  52. Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
  53. Rowley, J.J. & Cao, T.T. (2009) A new species of Leptolalax (Anura: Megophryidae) from central Vietnam. Zootaxa, 2198, 51–60.
  54. Rowley, J.J., Dau, V.Q. & Cao, T.T. (2017a) A new species of Leptolalax (Anura: Megophryidae) from Vietnam. Zootaxa, 4273 (1), 61–79. https://doi.org/10.11646/zootaxa.4273.1.5
  55. Rowley, J.J., Dau, V.Q., Hoang, H., Le, D.T., Cutajar, T.P. & Nguyen, T. (2017b) A new species of Leptolalax (Anura: Megophryidae) from northern Vietnam. Zootaxa, 4243 (3), 544–564. https://doi.org/10.11646/zootaxa.4243.3.7
  56. Rowley, J.J., Dau, Q.V. & Nguyen, T.T. (2013) A new species of Leptolalax (Anura: Megophryidae) from the highest mountain in Indochina. Zootaxa, 3737 (4), 415–428. https://doi.org/10.11646/zootaxa.3737.4.5
  57. Rowley, J.J., Hoang, H.D., Dau, V.Q., Le, D.T. & Cao, T.T. (2012) A new species of Leptolalax (Anura: Megophryidae) from central Vietnam. Zootaxa, 3321 (1), 56–68. https://doi.org/10.11646/zootaxa.3321.1.4
  58. Rowley, J.J., Hoang, H.D., Le, D.T.T., Dau, V.Q. & Cao, T.T. (2010a) A new species of Leptolalax (Anura: Megophryidae) from Vietnam and further information on Leptolalax tuberosus. Zootaxa, 2660 (1), 33–45. https://doi.org/10.11646/zootaxa.2660.1.3
  59. Rowley, J.J., Le, D.T.T., Tran, D.T.A. & Hoang, H.D. (2011) A new species of Leptolalax (Anura: Megophryidae) from southern Vietnam. Zootaxa, 2796 (1), 15–28. https://doi.org/10.11646/zootaxa.2796.1.2
  60. Rowley, J.J., Stuart, B.L., Neang, T. & Emmett, D.A. (2010b) A new species of Leptolalax (Anura: Megophryidae) from northeastern Cambodia. Zootaxa, 2567 (1), 57–68.
  61. Rowley, J.J., Stuart, B.L., Neang, T., Hoang, H.D., Dau, V.Q. & Nguyen, T.T. (2015a) A new species of Leptolalax (Anura: Megpphryidae) from Vietnam and Cambodia. Zootaxa, 4039 (3), 401–417. https://doi.org/10.11646/zootaxa.4039.3.1
  62. Rowley, J.J., Stuart, B.L., Richards, S.J., Phimmachak, S. & Sivongxay, N. (2010c) A new species of Leptolalax (Anura: Megophryidae) from Laos. Zootaxa, 2681 (1), 35–46. https://doi.org/10.11646/zootaxa.2681.1.3
  63. Rowley, J.J., Tran, D.T.A., Le, D.T.T., Dau, V.Q., Peloso, P.L.V., Nguyen, T.Q., Hoang, H.D., Nguyen, T.T. & Ziegler, T. (2016) Five new, microendemic Asian Leaf-litter Frogs (Leptolalax) from the southern Annamite mountains, Vietnam. Zootaxa, 4085 (1), 63−102. https://doi.org/10.11646/zootaxa.4085.1.3
  64. Sengupta, S., Sailo, S., Lalremsanga, H.T., Das, A. & Das, I. (2010) A new species of Leptolalax (Anura: Megophryidae) from Mizoram, northeastern India. Zootaxa, 2406 (1), 57–68. https://doi.org/10.11646/zootaxa.2406.1.3
  65. Shi, S., Hou, Y., Song, Z., Jiang, J.P. & Wang, B. (2021) A new leaf litter toad of Leptobrachella Smith, 1925 (Anura, Megophryidae) from Sichuan Province, China with supplementary description of L. oshanensis. Asian Herpetological Research, 12, 143–166. https://doi.org/10.16373/j.cnki.ahr.200118
  66. Simmons, J.E. (2002) Herpetological collecting and collections management. Revised edition. Society for the Study of Amphibians and Reptiles. Herpetological Circular, 31, 1–153.
  67. 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
  68. Smith, M.A. (1925) Contributions to the herpetology of Borneo. Sarawak Museum Journal, 3, 15–34.
  69. Stuart, B.L. & Rowley, J.J.L. (2020) A new Leptobrachella (Anura: Megophryidae) from the Cardamom Mountains of Cambodia. Zootaxa, 4834 (4), 556–572. https://doi.org/10.11646/zootaxa.4834.4.4
  70. Sung, Y.H., Yang, J. & Wang, Y. (2014) A new species of Leptolalax (Anura: Megophryidae) from southern China. Asian Herpetological Research, 5, 80–90. https://doi.org/10.3724/SP.J.1245.2014.00080
  71. Tamura, K. (1992) Estimation of the number of nucleotide substitutions when there are strong transition–transversion and G + C-content biases. Molecular Biology and Evolution, 9, 678–687.
  72. Tamura, K., Stecher, G. & Kumar, S. (2021). MEGA 11: Molecular Evolutionary Genetics Analysis Version 11. Molecular Biology and Evolution, 38 (7), 3022–3027. https://doi.org/10.1093/molbev/msab120.
  73. Tanabe, A.S. (2011) Kakusan 4 and Aminosan: two programs for comparing nonpartitioned, proportional and separate models for combined molecular phylogenetic analyses of multilocus sequence data. Molecular Ecology Resources, 11, 914–921. https://doi.org/10.1111/j.1755-0998.2011.03021.x
  74. Tavaré, S. (1986) Some probabilistic and statistical problems in the analysis of DNA sequences. Lectures on Mathematics in the Life Sciences, 17, 57–86.
  75. Taylor, E.H. (1962) The amphibian fauna of Thailand. University of Kansas Science Bulletin, 43, 265–599. https://doi.org/10.5962/bhl.part.13347
  76. Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTAL_X windows interface: flexible 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
  77. Wang, J., Li, Y.L., Li, Y., Chen, H.H., Zeng, Y.J., Shen, J.M. & Wang, Y.Y. (2019) Morphology, molecular genetics, and acoustics reveal two new species of the genus Leptobrachella from northwestern Guizhou Province, China (Anura, Megophryidae). ZooKeys, 848, 119–154. https://doi.org/10.3897/zookeys.848.29181
  78. Wang, J., Lyu, Z.T., Qi, S., Zeng, Z.C., Zhang, W.X., Lu, L.S. & Wang, Y.Y. (2020) Two new Leptobrachella species (Anura, Megophryidae) from the Yunnan-Guizhou Plateau, southwestern China. ZooKeys, 995, 97–125. https://doi.org/10.3897/zookeys.955.55939
  79. Wang, J., Qi, S., Dai, K.Y., Lyu, Z.T., Zeng, Z.C., Chen, H.H., Li, Y.Q., Zhao, Y.Y., Wang, Y.Z. & Wang, Y.Y. (2022) A new Leptobrachella species (Anura, Megophryidae) from South China, with comments on the taxonomic status of L. chishuiensis and L. purpurus. Zoosystematics and Evolution, 98, 165–180. https://doi.org/10.3897/zse.98.73162
  80. Wang, J., Yang, J.H., Li, Y., Lyu, Z.T., Zeng, Z.C., Liu, Z.Y., Ye, Y.H. & Wang, Y.Y. (2018) Morphology and molecular genetics reveal two new Leptobrachella species in southern China (Anura, Megophryidae). ZooKeys, 776, 105–137. https://doi.org/10.3897/zookeys.776.22925
  81. Yang, J., Wang, Y., Chen, G. & Rao, D.Q. (2016) A new species of the genus Leptolalax (Anura: Megophryidae) from Mt. Gaoligongshan of western Yunnan Province, China. Zootaxa, 4088, 379–394. https://doi.org/10.11646/zootaxa.4088.3.4
  82. Yang, J., Zeng, Z.C. & Wang, Y.Y. (2018) Description of two new sympatric species of the genus Leptolalax (Anura: Megophryidae) from western Yunnan of China. PeerJ, 6 (e4586), 1–32. https://doi.org/10.7717/peerj.4586
  83. Yuan, Z.-Y., Sun, R., Chen, J., Rowley, J.J.L., Wu, Z., Hou, S., Wang, S. & Che, J. (2017) A new species of the genus Leptolalax (Anura: Megophryidae) from Guangxi, China. Zootaxa, 4300(4), 551–570. https://doi.org/10.11646/zootaxa.4300.4.5