Abstract
Since its description from the Eaglenest Wildlife Sanctuary in Arunachal Pradesh of India, Leptobrachium bompu Sondhi & Ohler, 2011 has been considered a rarely sighted species with presumed low population density. The species continues to be known from a handful of specimens and little has been studied about it thereafter. Notably, the molecular identity of L. bompu from the type locality itself remains unassessed. Through our recent field surveys in the state, we have been able to study and dispel the misconceptions around the rarity of this species by reporting several new individuals here, alongside new locality records. In addition, we assess and establish the molecular identity of the typical and other reported populations of L. bompu from India for the first time, and clarify their phylogenetic relationships, both with closely related populations from nearby regions and all other 38 currently recognised members of the genus. Our various analyses confirm that the reported Chinese populations from Medog county, Tibet, previously referred to as L. ‘bompu’(herein L. cf. bompu), are divergent from L. bompu sensu stricto of India by genetic distances of 3.6–4.2% for the analysed fragment of mitochondrial DNA, suggesting that they could represent another potential cryptic species requiring further taxonomic scrutiny. Furthermore, our study employed an integrative approach to better understand the systematic affinities and biology of the poorly-known L. bompu. Based on our newly gathered data from the Indian populations, we provide a redescription of adult morphology, tadpole morphology, and vocal repertoire of male advertisement calls of the species, including comparative insights with respect to other previously documented species of the genus. Our work, therefore, not only improves the current knowledge on L. bompu, but also has resultant implications on the identity of related Chinese populations, alongside some genus-wide insights on Leptobrachium frogs at large.
References
- Al-Razi, H., Maria, M. & Poyarkov, N.A. (2021) Integrative taxonomic analysis reveals a new species of Leptobrachium Tschudi, 1838 (Anura, Megophryidae) from Bangladesh. Journal of Natural History, 55 (1–2), 85–114. https://doi.org/10.1080/00222933.2021.1894365
- Altig, R. (2007) A primer for the morphology of anuran tadpoles. Herpetological Conservation and Biology, 2, 71–74.
- Altig, R. & McDiarmid, R.W. (1999) Body plan. Development and morphology. In: McDiarmid, R.W. & Altig, R. (Eds.), Tadpoles: The Biology of Anuran Larvae. The University of Chicago Press, Chicago and London, pp. 24–51.
- AmphibiaWeb (2024) Amphibiaweb: information on amphibian biology and conservation. Electronic Database. University of California, Berkeley, California. Available from: https://amphibiaweb.org/ (accessed 10 October 2024)
- Bailey, V. (1933) The importance of types and type localities. Journal of Mammalogy, 14 (3), 241–243. https://doi.org/10.2307/1373822
- Bee, M.A., Suyesh, R. & Biju, S.D. (2013) Vocal behavior of the Ponmudi Bush Frog (Raorchestes graminirupes): repertoire and individual variation. Herpetologica, 69 (1), 22–35. https://doi.org/10.1655/HERPETOLOGICA-D-11-00042
- Biju, S.D., Garg, S., Mahony, S., Wijayathilaka, N., Senevirathne, G. & Meegaskumbura, M. (2014a) DNA barcoding, phylogeny and systematics of Golden-backed frogs (Hylarana, Ranidae) of the Western Ghats-Sri Lanka biodiversity hotspot, with the description of seven new species. Contributions to Zoology, 83 (4), 269–335. https://doi.org/10.1163/18759866-08304004
- Biju, S.D., Garg, S., Gururaja, K.V., Shouche, Y. & Walujkar, S.A. (2014b) DNA barcoding reveals unprecedented diversity in Dancing Frogs of India (Micrixalidae, Micrixalus): a taxonomic revision with description of 14 new species. Ceylon Journal of Science (Biological Sciences), 43, 37–123. https://doi.org/10.4038/cjsbs.v43i1.6850
- Biju, S.D., Garg, S., Gokulakrishnan, G., Chandrakasan, S., Thammachoti, P., Ren, J., Gopika, C., Bisht, K., Hamidy, A. & Shouche, Y. (2020) New insights on the systematics and reproductive behaviour in tree frogs of the genus Feihyla, with description of a new related genus from Asia (Anura, Rhacophoridae). Zootaxa, 4878 (1), 1–55. https://doi.org/10.11646/zootaxa.4878.1.1
- Bisht, K., Garg, S., Sarmah, A.N.D.A., Sengupta, S. & Biju, S.D. (2021) Lost, forgotten, and overlooked: systematic reassessment of two lesser-known toad species (Anura, Bufonidae) from Peninsular India and another wide-ranging northern species. Zoosystematics and Evolution, 97 (2), 451–470. https://doi.org/10.3897/zse.97.61770
- Brown, R.M., Siler, C.D., Diesmos, A.C. & Alcala, A.C. (2009) Philippine frogs of the genus Leptobrachium (Anura; Megophryidae): phylogeny-based species delimitation, taxonomic review, and descriptions of three new species. Herpetological Monograph, 23 (1), 1–44. https://doi.org/10.1655/09-037.1
- Charif, R.A., Strickman, L.M. & Waack, A.M. (2010) Raven Pro 1.4 User’s Manual. Cornell Laboratory of Ornithology, Ithaca, New York, 379 pp.
- Chen, W., Zhang, W., Zhou, S., Li, N., Huang, Y. & Mo, Y. (2013) Insight into the validity of Leptobrachium guangxiense (Anura: Megophryidae): evidence from mitochondrial DNA sequences and morphological characters. Zootaxa, 3641 (1), 31–40. https://doi.org/10.11646/zootaxa.3641.1.3
- Cochran, D.M. (1926) A new pelobatid batrachian from Borneo. Journal of the Washington Academy of Sciences, 16 (16), 446–447.
- Das, A., Garg, S., Hamidy, A., Smith, E.N. & Biju, S.D. (2019) A new species of Micryletta frog (Microhylidae) from Northeast India. PeerJ, 7, e7012. https://doi.org/10.7717/peerj.7012
- Delorme, M., Dubois, A., Grosjean, S. & Ohler, M. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24, 6–21.
- Ding, G.H., Chen, Z.Q., Tang, Y., Zheng, W.C. & Ji, X. (2020) The advertisement call of the moustache toad Leptobrachium liui Pope, 1947 (Anura: Megophryidae) from eastern China. Zootaxa, 4732 (4), 585–588. https://doi.org/10.11646/zootaxa.4732.4.8
- Draškić, G., Wangkulangkul, S., Martínez-Solano, I. & Vörös, J. (2018) Strong genetic subdivision in Leptobrachium hendricksoni (Anura: Megophryidae) in Southeast Asia. Amphibia-Reptilia, 39 (1), 99–111. https://doi.org/10.1163/15685381-17000102
- Dubois, A. & Ohler, A. (1998) A new species of Leptobrachium (Vibrissaphora) from northern Vietnam, with a review of the taxonomy of the genus Leptobrachium (Pelobatidae, Megophryinae). Dumerilia, 4, 1–32.
- Duellman, W.E. & Trueb, L. (1994) Biology of Amphibians. New York, McGraw-Hill, 670 pp.
- Eprilurahman, R. & Maghfiroh, N.L. (2020) The tadpole of Leptobrachium hasseltii Tschudi. 1838 (Amphibia: Anura: Megophryidae): Morphometry and larval developmental stage as identification character of species. AIP Conference Proceedings, 2260, 020023. https://doi.org/10.1063/5.0015681
- Fei, L., Hu, S.Q., Ye, C.Y. & Huang, Y.Z. (2009) Fauna Sinica. Amphibia. Vol. 2 Anura. Science Press, Beijing, 957 pp. [in Chinese]
- Fei, L., Ye, C.Y., Jiang, J.P., Cheng, S., Cai, C., Tang, Z., Cai, M., Gao, J. & Wei, G. (1995) Study on the classification and phylogenetic relationship of Vibrissaphora (Amphibia, Pelobatidae). Acta Herpetologica Sinica, 4–5, 65–71.
- Fei, L., Ye, C.Y., Jiang, J.P., Xie, F. & Huang, Y.Z. (2005) An Illustrated Key to Chinese Amphibians. Sichuan Publishing Group, Sichuan Publishing House of Science and Technology, Chengdu, 340 pp. [in Chinese]
- Fei, L., Ye, C.Y. & Jiang, J.P. (2010) Colored Atlas of Chinese Amphibians and Their Distributions. Sichuan Publishing House of Science and Technology, Chengdu, 519 pp. [in Chinese]
- Fischer, J.G. (1885) Ueber eine Kollektion von Amphibien und Reptilien aus Südost-Borneo. Archiv für Naturgeschichte, 51, 41–72. https://doi.org/10.5962/bhl.part.1568
- Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de S, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P.E., 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, 1–370. https://doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2
- Frost, D.R. (2024) Amphibian Species of the World: an online reference. version 6.1. Electronic Database. American Museum of Natural history, New York, New York. Available from: https://amphibiansoftheworld.amnh.org (accessed 10 October 2024)
- Fu, J., Weadick, C.J. & Bi, K. (2007) A phylogeny of the high- elevation Tibetan megophryid frogs and evidence for the multiple origins of reversed sexual size dimorphism. Journal of Zoology, 273 (3), 315–325. https://doi.org/10.1111/j.1469-7998.2007.00330.x
- Garg, S. & Biju, S.D. (2016) Molecular and morphological study of leaping frogs (Anura, Ranixalidae) with description of two new species. PloS One, 11 (11), e0166326. https://doi.org/10.1371/journal.pone.0166326
- Garg, S. & Biju, S.D. (2017) Description of four new species of Burrowing Frogs in the Fejervarya rufescens complex (Dicroglossidae) with notes on morphological affinities of Fejervarya species in the Western Ghats. Zootaxa, 4277 (4), 451–490. https://doi.org/10.11646/zootaxa.4277.4.1
- Garg, S. & Biju, S.D. (2019) New microhylid frog genus from Peninsular India with Southeast Asian affinity suggests multiple Cenozoic biotic exchanges between India and Eurasia. Scientific reports, 9 (1), 1906. https://doi.org/10.1038/s41598-018-38133-x
- Garg, S., Suyesh, R., Das, S., Bee, M.A. & Biju, S.D. (2021) An integrative approach to infer systematic relationships and define species groups in the shrub frog genus Raorchestes, with description of five new species from the Western Ghats, India. PeerJ, 9, e10791. https://doi.org/10.7717/peerj.10791
- 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
- Gopika, C., Garg, S., Sivaperuman, C., Gokulakrishnan, G. & Biju, S.D. (2023) Larval morphology and natural history in two Minervarya species from Andaman Islands, with comments on a new phytotelmonous larval morphotype for the genus. Zootaxa, 5353 (6), 567–581. https://doi.org/10.11646/zootaxa.5353.6.4
- Gosner, K.L. (1960) A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16, 183–190.
- Grosjean, S. (2001) The tadpole of Leptobrachium (Vibrissaphora) echinatum (Amphibia, Anura, Megophryidae). Zoosystema, 23, 143–156.
- Haas, A., Das, I., Hertwig, S.T., Bublies, P. & Schulz-Schaeffer, R. (2022) A Guide to the Tadpoles of Borneo (eBook version). Tredition, Hamburg, Germany, 553 pp.
- Hamidy, A. & Matsui, M. (2010) A new species of blue-eyed Leptobrachium (Anura: Megophryidae) from Sumatra, Indonesia. Zootaxa, 2395 (1), 34–44. https://doi.org/10.11646/zootaxa.2395.1.3
- Hamidy, A. & Matsui, M. (2014) A new species of Leptobrachium from the Kelabit highland, Northwestern Borneo (Anura, Megophryidae). Current Herpetology, 33 (1), 57–67. https://doi.org/10.5358/hsj.33.57
- Hamidy, A. & Matsui, M. (2017) Phylogenetic relationships of Leptobrachium hasseltii Tschudi, 1838 (Amphibia, Anura, Megophryidae) - Detection of a possible cryptic species. Treubia, 44, 15–28. https://doi.org/10.1016/j.ympev.2010.03.014
- Hamidy, A., Matsui, M., Shimada, T., Nishikawa, K., Yambun, P., Sudin, A., Kusrini, M.D. & Kurniati, H. (2011) Morphological and genetic discordance in two species of Bornean Leptobrachium (Amphibia, Anura, Megophryidae). Molecular Phylogenetics and Evolution, 61 (3), 904–913. https://doi.org/10.1016/j.ympev.2011.08.020
- Hamidy, A., Matsui, M., Nishikawa, K. & Belabut, D.M. (2012) Detection of cryptic taxa in Leptobrachium nigrops (Amphibia, Anura, Megophryidae), with description of two new species. Zootaxa, 3398 (1), 22–39. https://doi.org/10.11646/zootaxa.3398.1.2
- 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
- Hu, S.C., Djao, E.M. & Liu, C.C. (1973) A survey of Amphibians and Reptiles in Kweichow province, including a herpetofaunal analysis. Acta Zoologica Sinica, 19 (2), 149–178. [in Chinese]
- Huang, X.Y., Zhang, X.C., Yang, S.K. & Yang, J.H. (2019) Acoustic characteristics of male calls of Leptobrachium tengchongense in breeding season. Chinese Journal of Zoology, 54, 189–195.
- Huelsenbeck, J.P., Ronquist, F., Neilsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294 (5550), 2310–2314. https://doi.org/10.1126/science.1065889
- IUCN. (2022) The IUCN Red List of Threatened Species. Version 2022-2. Available from: https://www.iucnredlist.org/ (accessed 10 October 2024)
- K. Lisa Yang Center for Conservation Bioacoustics at the Cornell Lab of Ornithology (2024) Raven Pro: Interactive Sound Analysis Software Version 1.6.4. The Cornell Lab of Ornithology, Ithaca, New York. Available from: https://ravensoundsoftware.com/ [Computer software]
- Köhler, J., Jansen, M., Rodriguez, A., Kok, P.J., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F., Roedel, M.O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251 (1), 1–124. https://doi.org/10.11646/zootaxa.4251.1.1
- Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, 33 (7), 1870–1874. https://doi.org/10.1093/molbev/msw054
- Lalremsanga, H.T., Sailo, S. & Zonunsanga, H.T. (2009) “Quack quack quack”-revealing the strange advertisement call of Leptobrachium smithi (Anura: Pelobatidae): a bioacoustic analysis. Science Vision, 9, 121–129.
- 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, 772–773. https://doi.org/10.1093/molbev/msw260
- Liu, C.C. (1945) New frogs from West China. Journal of the West China Border Research Society, Series B, 15, 28–44.
- Liu, C.C. & Hu, S.C. (1962) A herpetological report of Kwangsi. Acta Zoologica Sinica, 14, 73–104. [in Chinese]
- Lê, D.T.T., Tran, L.T.C., Huynh, A.V.H. & Tran, D.T.A. (2018) Tadpole morphology of the white-eyed toad Leptobrachium leucops Stuart, Rowley, Tran, Le & Hoang 2011 (Anura: Megophryidae) in Bidoup-Nui Ba National Park, Lam Dong province. Science & Technology Development Journal: Natural Sciences, 2 (2), 14–21. https://doi.org/10.32508/stdjns.v2i2.729
- Leaché, A.D. & Reeder, T.W. (2002) Molecular systematics of the Eastern Fence Lizard (Sceloporus undulatus): A comparison of parsimony, likelihood, and bayesian approaches. Systematics Biology, 51 (1), 44–68. https://doi.org/10.1080/106351502753475871
- Liang, X., Liu, W., Wang, B., Ding, L., Wu, J., Xie, F. & Jiang, J. (2017) Leptobrachium bompu (Amphibia, Anura, Megophryidae) discovered in Upper Medog, Tibet, China with descriptions of its tadpoles, advertisement calls and systematic position. Asian Herpetological Research, 8 (2), 137–146. https://doi.org/10.16373/j.cnki.ahr.160023
- Liu, C.C. (1945) New frogs from west China. Journal of West China Border Research Society, Series B, 15, 28–43.
- Malkmus, R. (1996) Leptobrachium gunungensis sp. n. (Anura: Pelobatidae) vom Mount Kinabalu, Nord‐Borneo. Mitteilungen aus dem Museum für Naturkunde in Berlin. Zoologisches Museum und Institut für Spezielle Zoologie, Berlin, 72 (2), 297–301. https://doi.org/10.1002/mmnz.19960720209
- Manthey, U. & Denzer, W. (2013) Südostasiatische Anuren im Fokus, Spezies der Gattung Leptobrachium Tschudi, 1838 (Amphibia, Anura, Megophryidae). Sauria, 35 (3), 3–22.
- Matsui, M. (2013) A new Leptobrachium (Vibrissaphora) from Laos (Anura: Megophryidae). Current Herpetology, 32 (2), 182–189. https://doi.org/10.5358/hsj.32.182
- Matsui, M., Nabhitabhata, J. & Panha, S. (1999) On Leptobrachium from Thailand with a description of a new species (Anura: Pelobatidae). Japanese Journal of Herpetology, 18, 19–29. https://doi.org/10.5358/hsj1972.18.1_19
- 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. https://doi.org/10.1016/j.ympev.2010.03.014
- 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
- Minh, B.Q., Nguyen, M.A.T. & Haeseler, A.V. (2013) Ultra-fast approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30 (5), 1188–1195. https://doi.org/10.1093/molbev/mst024
- Myers, C.W. & Duellman, W.E. (1982) A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from Western Panama. American Museum Novitates, 2752, 1–32.
- Nguyen, L.T., Schmidt, H.A., Von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32 (1), 268–274. https://doi.org/10.1093/molbev/msu300
- Pawangkhanant, P., Poyarkov, N.A., Duong, T.V., Naiduangchan, M. & Suwannapoom, C.A. (2018) New species of Leptobrachium (Anura, Megophryidae) from western Thailand. PeerJ, 6 (6), e5584. https://doi.org/10.7717/peerj.5584
- Pham, V.A., Le, D.T., Pham, C.T., Nguyen, S.L.H., Ziegler, T. & Nguyen, T.Q. (2016) Two additional records of megophryid frogs, Leptobrachium masatakasatoi Matsui, 2013 and Leptolalax minimus (Taylor, 1962), for the herpetofauna of Vietnam. Revue suisse de Zoologie, 123 (1), 35–43.
- Pope, C.H. (1947) A new pelobatid frog from Fukien Province, China. Copeia, 1947, 109–112. https://doi.org/10.2307/1438642
- Rao, D.Q. & Wilkinson, J.A. (2008) Phylogenetic relationships of the mustache toads inferred from mtDNA sequences. Molecular Phylogenetics and Evolution, 46 (1), 61–73. https://doi.org/10.1016/j.ympev.2007.10.005
- Rao, D.Q., Wilkinson, A. & Zhang, M.W. (2006) A new species of the genus Vibrissaphora (Anura: Megophryidae) from Yunnan Province, China. Herpetologica, 62 (1), 90–108. https://doi.org/10.1655/05-05.1
- Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
- Saikia, B., Sinha, B. & Kharkongor, I.J. (2017) A second record of the Eastern Spadefoot Toad (Amphibia: Anura: Megophryidae: Leptobrachium bompu Sondhi & Ohler, 2011) with a note on its morphological variations and natural history. Journal of Threatened Taxa, 9 (9), 10692–10696. https://doi.org/10.11609/jott.3300.9.9.10692-10696
- Savage, J.M. & Heyer, W.R. (1967) Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America: With 6 figures. Studies on Neotropical Fauna and Environment, 5, 111–131. https://doi.org/10.1080/01650526709360400
- 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 Entomo- logical Society of America, 87 (6), 651–701. https://doi.org/10.1093/aesa/87.6.651
- Singh, P., Dey, M. & Ramanujam, S.N. (2016) A description of the tadpole of Leptobrachium smithi (Anura: Megophryidae) from South Assam, India. Herpetological Conservation and Biology, 11 (2), 280–285.
- Sondhi, S. & Ohler, A. (2011) A blue-eyed Leptobrachium (Anura: Megophryidae) from Arunachal Pradesh, India. Zootaxa, 2912 (1), 28–36. https://doi.org/10.11646/zootaxa.2912.1.2
- Stuart, B.L., Sok, K. & Neang, T. (2006) A collection of amphibians and reptiles from hilly eastern Cambodia. The Raffles Bulletin of Zoology, 54 (1), 129–155.
- Stuart, B.L., Rowley, J.J.L., Tran, D.T.A., Le, D.T.T. & Hoang, H.D. (2011) The Leptobrachium (Anura: Megophryidae) of the Langbian Plateau, southern Vietnam, with description of a new species. Zootaxa, 2804, 25–40. https://doi.org/10.11646/zootaxa.2804.1.3
- Stuart, B.L., Phimmachak, S., Seateun, S. & Sivongxay, N. (2012) A new Leptobrachium (Anura: Megophryidae) from the highlands of southeastern Laos. Zootaxa, 3155 (1), 29–37. https://doi.org/10.11646/zootaxa.3155.1.3
- Stuart, B.L., Som, H.E., Neang, T., Hoang, H.D., Le, D.T.T., Dau, V.Q., Potter, K. & Rowley, J.J. (2020) Integrative taxonomic analysis reveals a new species of Leptobrachium (Anura: Megophryidae) from north-eastern Cambodia and central Vietnam. Journal of Natural History, 54 (1–4), 225–255. https://doi.org/10.1080/00222933.2020.1756498
- Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b10. Sinauer Association Inc., Sunderland, Massachusetts. [program]
- Tenzin, J. & Wangyal, J.T. (2019) New record of Blue-eyed Eastern Spadefoot Toad Leptobrachium bompu (Amphibia: Megophryidae) from Sarpang District in Bhutan. Journal of Threatened Taxa, 11 (3), 13385–13389. https://doi.org/10.11609/jott.4134.11.3.13385-13389
- Thomas, A., Suyesh, R., Biju, S.D. & Bee, M.A. (2014) Vocal behavior of the elusive purple frog of India (Nasikabatrachus sahyadrensis), a fossorial species endemic to the Western Ghats. PLoS One, 9 (2), e84809. https://doi.org/10.1371/journal.pone.0084809
- Thompson, J.D., Higgins, D.G. & Gibson, T.J. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research, 22 (22), 4673–4680. https://doi.org/10.1093/nar/22.22.4673
- Tschudi, J.J.von. (1838) Classification der Batrachier mit Berücksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Petitpierre, Neuchâtel, 99 pp. https://doi.org/10.5962/bhl.title.4883
- Wogan, G.O.U. (2012) A new species of Leptobrachium from Myanmar (Anura: Megophryidae). Zootaxa, 3415 (1), 23–36. https://doi.org/10.11646/zootaxa.3415.1.2
- Xu, W., Dong, W.J., Fu, T.T., Gao, W., Lu, C.Q., Yan, F., Wu, Y.H., Jiang, K., Jin, J.Q., Chen, H.M. & Zhang, Y.P. (2021) Herpetological phylogeographic analyses support a Miocene focal point of Himalayan uplift and biological diversification. National Science Review, 8 (9), nwaa263. https://doi.org/10.1093/nsr/nwaa263
- Yang, D.T., Ma, D.S., Li, F.L. & Chen, H.J.J. (1983) Descriptions of two new pelobatid toads from Yunnan. Acta Zootaxonomica Sinica, 8, 323–327.
- Yang, J., Wang, Y. & Chan, B.P.L. (2016) A new species of the genus Leptobrachium (Anura: Megophryidae) from the Gaoligongshan Mountain Range, China. Zootaxa, 4150 (2), 133–148. https://doi.org/10.11646/zootaxa.4150.2.3
- Zheng, Y., Li, S. & Fu, J. (2008) A phylogenetic analysis of the frog genera Vibrissaphora and Leptobrachium, and the correlated evolution of nuptial spine and reversed sexual size dimorphism. Molecular Phylogenetics and Evolution, 46 (2), 695–707. https://doi.org/10.1016/j.ympev.2007.09.019
- Zheng, Y., Deng, D., Li, S. & Fu, J. (2010) Aspects of the breeding biology of the Omei mustache toad (Leptobrachium boringii): polygamy and paternal care. Amphibia-Reptilia, 31 (2), 183–194. https://doi.org/10.1163/156853810791069092
- Zheng, Y., Rao, D., Murphy, R.W. & Zeng, X. (2011) Reproductive behavior and underwater calls in the Emei mustache toad, Leptobrachium boringii. Asian Herpetological Research, 2 (4), 199–215. https://doi.org/10.3724/SP.J.1245.2011.00199