Abstract
The Megophrys major species group (MMSG) is composed of typically medium to large sized frogs. Within the genus, it is the most geographically widespread clade ranging from the western Himalayas to southern Indochina. In this study, we examined in detail the extent of cryptic diversity within the MMSG-Indian populations based on molecular data (up to ten genes) using multigene concatenation and coalescent-based phylogenetic techniques, species delimitation analyses and extensive morphological data.
Molecular evidence suggests a high level of hidden cryptic diversity within the MMSG, particularly within the M. major species complex (MMC), highlighting overlapping distributions, a case of potential mitochondrial transfer between two species, and tree topology discordance between phylogenetic methods and mitochondrial and nuclear data sets. Most analyses indicated distinct eastern and western clades in the MMC, and that the western clade may further divide into a northern and a southern subclade.
A detailed taxonomic review of Indian members of the Megophrys major species group is provided. Previously undocumented complex nomenclatural issues involving known species are highlighted and resolved. Megophrys monticola is taxonomically redefined for the first time as a valid species since its synonymy with M. parva in 1893. The taxonomic status of two recently described species, Xenophrys katabhako and X. sanu are discussed in light of increased molecular and morphological sampling, and are synonymised with M. monticola. Megophrys monticola and M. robusta are redescribed based on their original type specimens and recently collected material. Megophrys major is neotypified and M. robusta lectotypified to remove prevailing nomenclatural instability. Four new large sized Indian MMC species are formally described from the Northeast Indian states of Arunachal Pradesh, Meghalaya and Nagaland, and Myanmar. All South Asian MMSG species are morphologically diagnosed from known congeners in the group. The geographic distributions of all taxa discussed are significantly redefined based on the revised taxonomy and extensive literature review. Morphological and molecular evidence suggests that Megophrys major sensu stricto might be endemic to Northeast India; M. glandulosa is formally removed from the Indian and Bhutan species checklists. Numerous misidentifications in literature are highlighted and corrected. In order to reduce future misidentifications of species reported from surrounding regions, high definition images of the holotypes of three Chinese species, M. glandulosa, M. medogensis and M. zhangi are provided for the first time, and a detailed description of Myanmar specimens of M. glandulosa is also given. This study provides the principle foundation for further research into the taxonomic status of the remaining, currently undescribed MMC taxa from Southeast Asia.
References
Ahmed, A.F., Das, A. & Dutta, S.K. (2009) Amphibians and Reptiles of Northeast India. A Photographic Guide. Aaranyak, Guwahati, 169 pp.
Anders, C. (2002) Class Amphibia (amphibians). In: Schleich, H.H. & Kästle, W. (Eds.), Amphibians and Reptiles of Nepal. A.R.G. Gantner Verlag K.G., Ruggell, pp. 133–340.
Anderson, J. (1871a) 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.
Anderson, J. (1871b) On some Indian reptiles. Proceedings of the Zoological Society of London, 1871, 149–211.
Anderson, J. (1879 "1878") Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the Two Expeditions to Western Yunnan in 1868 and 1875; and a Monograph of the Two Cetacean Genera Platanista and Orcella. Vol. 1 & 2. Bernard Quaritch, London, XXV + 985 pp. & 81 pls.
Annandale, N. (1906) Notes on the freshwater fauna of India. No. XIII. Some Himalayan tadpoles. Journal of the Asiatic Society of Bengal, 2 (7), 289–292.
Annandale, N. (1911) Contributions to the fauna of Yunnan based on collections made by J. Coggin Brown, B.Sc., 1909–1910. Records of the Indian Museum, 6, 215–218.
Annandale, N. (1912) Zoological results of the Abor Expedition, 1911–1912. I. Batrachia. Records of the Indian Museum, 8 (1), 7–36, pls. II–IV.
https://doi.org/10.5962/bhl.part.1186Anonymous (1994) Opinion 1763. Megophrys montana Kuhl & Van Hasselt, 1822 (Amphibia, Anura): generic and specific names placed on Official Lists, and Leptobrachium parvum Boulenger, 1893 (currently Megophrys parva): specific name conserved. Bulletin of Zoological Nomenclature, 51, 84–85.
Ao, M., Bordoloi, S. & Ohler, A. (2003) Amphibian fauna of Nagaland with nineteen new records from the State including five new records for India. Zoos' Print Journal, 18 (6), 1117–1125.
https://doi.org/10.11609/JoTT.ZPJ.18.6.1117-25Bauer, A.M., Parham, J.F., Brown, R.M., Stuart, B.L., Grismer, L., Papenfuss, T.J., Wolfgang, B., Savage, J.M., Carranza, S., Grismer, J.L., Wagner, P., Schmitz, A., Ananjeva, N.B. & Inger, R.F. (2010) Availability of new Bayesian-delimited gecko names and the importance of character-based species descriptions. Proceedings of the Royal Society of London B: Biological Sciences, 278, 490–492.
https://doi.org/10.1098/rspb.2010.1330Bain, R.H. & Nguyen, Q.T. (2004) Herpetofaunal diversity of Ha Giang Province in northeastern Vietnam, with descriptions of two new species. Novitates, 3453, 1–42.
https://doi.org/10.1206/0003-0082(2004)453<0001:HDOHGP>2.0.CO;2
Bain, R.H., Lathrop, A., Murphy, R.W., Orlov, N.L. & Cuc, H.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<0001:CSOACF>2.0.CO;2Benson, D.A., Cavanaugh, M., Clark, K., Karsch-Mizrachi, I., Lipman, D.J., Ostell, J. & Sayers, E.W. (2017) GenBank. Nucleic Acids Research, 46 (D1), 41–47. [D41–D47]
https://doi.org/10.1093/nar/gkw1070Biju, S.D. & Bossuyt, F. (2003) New frog family from India reveals an ancient biogeographical link with the Seychelles. Nature, 425, 711–714.
https://doi.org/10.1038/nature02019Biju, S.D., 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, 31–46.
Biju, S.D., Garg, S., Mahony, S., Wijayathilaka, N., Senevirathne, G. & Meegaskumbura, M. (2014) 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, 269–335.
Biju, S.D., Senevirathne, G., Garg, S., Mahony, S., Kamei, R.G., Thomas, A., Shouche, Y., Raxworthy, C.J., Meegaskumbura, M. & van Bocxlaer, I. (2016) Frankixalus, a new rhacophorid genus of tree hole breeding frogs with oophagous tadpoles. PLOS ONE, 11 (1), e0145727.
https://doi.org/10.1371/journal.pone.0145727Blyth, E. (1855 [“1854”]) Notices and descriptions of various reptiles, new or little known. Journal of the Asiatic Society of Bengal, 23, 287–302.
Borah, M.M. & Bordoloi, S. (2001) Nine new records of Amphibia (Anura) from Arunachal Pradesh, India. Journal, Bombay Natural History Society, 98 (1), 123–126.
Bordoloi, S. & Borah, M.M. (2001) First record of a frog Megophrys robusta (Boulenger 1908) from Arunachal Pradesh, India. Cotton College Research Journal, 1, 191–193.
Bordoloi, S., Borah, M.M., Sharma, K.K. & Kalita, S.N. (2000) An overview of amphibian fauna of Arunachal Pradesh, India. Journal of the Assam Science Society, 41 (4), 363–371.
Bouckaert, R.R. (2010) DensiTree: making sense of sets of phylogenetic trees. Bioinformatics, 26, 1372–1373.
https://doi.org/10.1093/bioinformatics/btq110Bouckaert, R., Alvarado-Mora, M. & Rebello Pinho, J.A. (2013) Evolutionary rate and HBV: issues of rate estimation with Bayesian molecular methods. Antiviral Therapy, 18, 497–503.
https://doi.org/10.3851/IMP2656Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Caudata in the Collections of the British Museum. (Natural History). Second Edition. Taylor and Francis, London, XVI + 503 pp., XXX pls.
Boulenger, G.A. (1886) Description of a new frog of the genus Megalophrys. Proceedings of the Zoological Society of London, 1885, 850, pl. 1.
Boulenger, G.A. (1889) Description of a new batrachian of the genus Leptobrachium, obtained by M.L. Fea in the Karens Mountains, Burma. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 7, 748–750.
Boulenger, G.A. (1890) The Fauna of British India, Including Ceylon and Burma. Reptilia and Batrachia. Taylor & Francis, London, XVIII + 541 pp.
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 1887–8. Annali del Museo Civico di Storia Naturale di Genova, Series 2, 13, 304–347, pls. VII–XII.
Boulenger, G.A. (1899) On a collection of reptiles and batrachians made by Mr. J.D. La Touche in N.W. Fokien, China. Proceedings of the Zoological Society of London, 1899, 159–172, pls. XVI–XIX.
Boulenger, G.A. (1903) Report on the batrachians and reptiles. In: Annandale, N. & Robinson H.C. (Eds.), Fasciculi Malayenses. Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States 1901–1903 Undertaken by Nelson Annandale and Herbert C. Robinson Under the Auspices of the University of Edinburgh and the University of Liverpool. Vol. 2. Zoology. Part 1. Longmans, Green & Co., London, pp. 131–176, pls. V–VI.
Boulenger, G.A., Annandale, N., Wall, F. & Regan, C.T. (1907) Reports on a collection of Batrachia, reptiles and fish from Nepal and the Western Himalayas. Records of the Indian Museum, 1, 149–158, pl. VI.
Boulenger, G.A. (1908) A revision of the oriental pelobatid batrachians (genus Megalophrys). Proceedings of the Zoological Society of London, 1908, 407–430, pls. XXII–XXV, fig. 71.
https://doi.org/10.1111/j.1096-3642.1908.tb01852.xBourret, R. (1937) Notes herpétologiques sur l’Indochine française. XIV. Les Batraciens de la collection du Laboratoire des Sciences Naturelles de l’Université. Descriptions de quinze espèces ou variétés nouvelles. Annexe au Bulletin Général de l'Instruction Publique, 4, 5–56.
Bourret, R. (1942) Les batrachians l’Indochine. Mémoires de l’Institut Océanographique de L’Indochine, 1–547 + I–X, pls. 1–4.
Carstens, B.C. & Knowles, L.L. (2007) Estimating species phylogeny from gene-tree probabilities despite incomplete lineage sorting: an example from Melanoplus grasshoppers. Systematic Biology, 56 (3), 400–411.
https://doi.org/10.1080/10635150701405560Chanda, S.K. (1986) On a collection of anuran amphibians from Darjeeling and Sikkim Himalayas, with description of a new species of Rana (Ranidae). Journal of Bengal Natural History Society, 5 (2), 140–155.
Chanda, S.K. (2002) Handbook - Indian Amphibians. Zoological Survey of India, Kolkata, 335 pp.
Chanda, S.K., Das, I & Dubois, A. (2000 “1999”) Catalogue of amphibian types in the collection of Zoological Survey of India. Hamadryad, 25 (2), 100–128.
Che, J., Chen, H.-M., Yang, J.-Z., Jin, J.-Q., Jiang, K., Yuan, Z.-Y., Murphy, R.W. & Zhang, Y.-P. (2012) Universal COI primers for DNA barcoding amphibians. Molecular Ecology Resources, 12 (2), 247–258.
https://doi.org/10.1111/j.1755-0998.2011.03090.xChen, J.-M., Zhou, W.-W., Poyarkov Junior, N.A. Stuart, B.L., Brown, R.M., Lathrop, A., Wang, Y.-Y., Yuan, Z.-Y., Jiang, K., Hou, M., Chen, H.-M., Suwannapoom, C., Ngoc Nguyen, S., Van Duong, T., Papenfuss, T.J., Murphy, R.W., Zhang, Y.-P. & Che, J. (2017) A novel multilocus phylogenetic estimation reveals unrecognized diversity in Asian horned toads, genus Megophrys sensu lato (Anura: Megophryidae). Molecular Phylogenetics and Evolution, 106, 28–43.
https://doi.org/10.1016/j.ympev.2016.09.004Cranston, K., Hurwitz, B., Ware, D., Stein, L. & Wing, R. (2009) Species trees from highly incongruent gene trees in rice. Systematic Biology, 58 (5), 489–500.
https://doi.org/10.1093/sysbio/syp054Daniel, J.C. (1962) Notes on some amphibians of the Darjeeling area, West Bengal. Journal of the Bombay Natural History Society, 59 (2), 666–668.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772.
https://doi.org/10.1038/nmeth.2109Das, I., Tron, R.K.L., Rangad, D. & Hooroo, R.N. (2010) A new species of Leptolalax (Anura: Megophryidae) from the sacred groves of Mawphlang, Meghalaya, north-eastern India. Zootaxa, 2339, 44–56.
Degnan, J.H. & Rosenberg, N.A. (2009) Gene tree discordance, phylogenetic inference and the multispecies coalescent. Trends in Ecology & Evolution, 24 (6), 332–340.
https://doi.org/10.1016/j.tree.2009.01.009
Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24 (1–4), 6–21.
Deuti, K & Ayyaswamy, A.K. (2008) Three new records of amphibians from West Bengal. Zoological Survey of India, 108 (4), 17–20.
Deuti, K & Kumar, A. (2008) Rediscovery of Robust pelobatid frog, Xenophrys robusta, after fifty years with three colour morphs. Zoological Survey of India, 108 (4), 119–120.
Deuti, K., Grosjean, S., Nicolas, V., Vasudevan, K. & Ohler, A. (2017) Nomenclatural puzzle in early Xenophrys nomina (Megophryidae, Amphibia) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34, 20–48.
Dever, J.A., Fuiten, A.M., Konu, Ö. & Wilkinson, J.A. (2012) Cryptic torrent frogs of Myanmar: An examination of the Amolops marmoratus species complex with the resurrection of Amolops afghanus and the identification of a new species. Copeia, 2012, 57–76.
https://doi.org/10.1643/CH-10-180Dubois, A. (1982) Le statut nomenclatural des noms génériques d’Amphibiens Anures créés par Kuhl & Van Hasselt (1822): Megophrys, Occidozyga et Rhacophorus. Bulletin du Museum National d'Histoire Naturelle, Paris, Series 3, 4A (1–2), 261–280.
Dubois, A. (1989 “1988”) Leptobrachium parvum Boulenger, 1893 (Amphibia, Anura): proposed conservation. Alytes, 7 (3), 97–100.
Dubois, A. (1992) Case 2382. Megophrys montana Kuhl & van Hasselt, 1822 (Amphibia, Anura): proposed placement of both the generic and specific names on Official Lists, and Leptobrachuim parvum Boulenger, 1893 (currently Megophrys parva): proposed conservation of the specific name. Bulletin of Zoological Nomenclature, 49 (3), 213–216
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), 1–32.
Dutta, S.K. (1997) Amphibians of India and Sri Lanka (Checklist and Bibliography). Odyssey Publishing House, Bubaneswar, XIII + 342 + XXII pp.
Fei, L. (1999) Atlas of Amphibians of China. Publishing House for Scientific and Technological Literature, Hunan, 432 pp.
Fei, L., Ye, C.Y., Wu, G.F. & Hu, S.Q. (1977) A survey of amphibians in Xizang (Tibet). Acta Zoologica Sinica, 23 (1), 54–63.
Fei, L., Ye, C.-Y. & Huang, Y.-Z. (1983) Two new subspecies of Megophrys omeimontis Liu from China (Amphibia; Pelobatidae). Acta Herpetologica Sinica, 2 (2), 49–52.
Fei, L., Ye, C.-Y. & Huang, Y.-Z. (1990) Key to Chinese Amphibians. Chongqing Branch, Science and Technology Literature Publishing House, Chongqing, 364 + 2 + [iv] pp.
Fei, L., Ye, C.-Y. & Huang, Y.-Z. (1992) Two new species of the genus Megophrys, Pelobatidae (Amphibia, Anura) from China. Zoological Research, 13 (1), 5–12.
Fei, L., Hu, S.-Q., Ye, C.-Y. & Huang, Y.-Z. (2009) Fauna Sinica. Amphibia. Volume 2. Anura. Chinese Academy of Science, Science, Beijing, XIII + 957 pp.
Fei, L., Ye, C. & Jiang, J. (2010) Coloured Atlas of Chinese Amphibians. Sichuan Publishing House of Science & Technology, Sichuan, 517 pp.
Fei, L., Ye, C.-Y. & Jiang, J.-P. (2012) Coloured Atlas of Chinese Amphibians and their Distributions. Sichuan Publishing House of Science & Technology, Sichuan, 620 pp.
Flot, J.F. (2010) Seqphase: a web tool for interconverting phase input/output files and fasta sequence alignments. Molecular Ecology Resources, 10 (1), 162–166.
https://doi.org/10.1111/j.1755-0998.2009.02732.xFrost, D.L., Grant, T., Faivovich, J.N., Bain, R.H., Haas, A., Haddad, C.F.B., De Sa´, R., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, R.C. (2006) 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;2Fu, C., Hua, X., Li, J., Chang, Z., Pu, Z. & Chen, J. (2006) Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects. Ecography, 29, 919–927 + 1–7 (Appendix E4802).
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.xFujita, M.K. & Leaché, A.D. (2010) A coalescent perspective on delimiting and naming species: a reply to Bauer et al. Royal Society of London B: Biological Sciences, 278, 493–495.
https://doi.org/10.1098/rspb.2010.1864Fujita, M.K., Leaché, A.D., Burbrink, F.T., McGuire, J.A. & Moritz, C. (2012) Coalescent-based species delimitation in an integrative taxonomy. Trends in Ecology & Evolution, 27 (9), 480–488.
https://doi.org/10.1016/j.tree.2012.04.012Fusinatto, L.A., Alexandrino, J., Haddad, C.F.B., Brunes, T.O., Rocha, C.F.D. & Sequeira, F. (2013) Cryptic genetic diversity is paramount in small-bodied amphibians of the genus Euparkerella (Anura: Craugastoridae) endemic to the Brazilian Atlantic Forest. PLOS ONE, 8 (11), e79504.
https://doi.org/10.1371/journal.pone.0079504García-París, M., Buchholz, D.R. & Parra-Olea, G. (2003) Phylogenetic relationships of Pelobatoidea re-examined using mtDNA. Molecular Phylogenetics and Evolution, 28 (1), 12–23.
https://doi.org/10.1016/S1055-7903(03)00036-8Günther, A. (1860) Contributions to a knowledge of the reptiles of the Himalaya mountains. I. Descriptions of the new species. II. List of Himalayan reptiles, with remarks on their horizontal distribution. Proceedings of the Zoological Society of London, 1860, 148–175.
Günther, A. (1864) The Reptiles of British India. The Ray Society, London, 452 + XXVII pp., 26 pls.
Günther, A. (1868) First account of species of tailless batrachians added to the collection of the British Museum. Proceedings of the Zoological Society of London, 1868, 478–490, pls. XXXVII–XL.
Günther, A. (1875) Second report on collections of Indian reptiles obtained by the British Museum. Proceedings of Zoological Society, 1875 (1), 224–234.
Günther, A. (1876) Third report on collections of Indian reptiles obtained by the British Museum. Proceedings of Zoological Society, 1875 (4), 567–577.
Hasan, M., Islam, M.M., Khan, M.M.R., Igawa, T., Alam, M.S., Djong, H.T., Kurniawan, N., Joshy, H., Sen, Y.H., Belabut, D.M. & Kurabayashi, A. (2014) Genetic divergences of South and Southeast Asian frogs: a case study of several taxa based on 16S ribosomal RNA gene data with notes on the generic name Fejervarya. Turkish Journal of Zoology, 38 (4), 389–411.
https://doi.org/10.3906/zoo-1308-36Heled, J. & Drummond, A.J. (2009) Bayesian inference of species trees from multilocus data. Molecular Biology and Evolution, 27 (3), 570–580.
https://doi.org/10.1093/molbev/msp274Hoegg, S., Vences, M., Brinkmann, H. & Meyer, A. (2004) Phylogeny and comparative substitution rates of frogs inferred from sequences of three nuclear genes. Molecular Biology and Evolution, 21 (7), 1188–1200.
https://doi.org/10.1093/molbev/msh081Hooker, J.D. (1854) Himalayan Journals: Notes of a Naturalist in Bengal, the Sikkim and Nepal Himalayas, the Khasia Mountains. Vol. II. John Murray, London, XII + 487 pp., pl. XII.
Hora, S.L. (1923) Observations on the fauna of certain torrential streams in the Khasi Hills. Records of the Indian Museum, 25, 579–599.
Hu, S.-C., Djao, E.-M. & Liu, C.-C. (1966) A herpetological survey of the Tsinling and Ta-Pa Shan region. Acta Zoologica Sinica, 18 (1), 57–89, pls. I–III.
Huang, C.-M., Lathrop, A. & Murphy, R.W. (1998) Translations of "Two new species of amphibian in Tibet" Huang & Fei 1981 and "Description of two new species of the genus Megophrys" Fei, Ye, & Huang 1995[1992]. Smithsonian Herpetological Information Service, 118, 1–20.
Huang Y-Z. & Fei, L. (1981) Two new species of amphibians from Tibet. Acta Zootaxonomica Sinica, 6 (2), 211–215.
Humtsoe, L.N., Bordoloi, S., Ohler, A. & Dubois, A. (2008) Rediscovery of a long known species, Ixalus lateralis Anderson, 1871. Zootaxa, 1921, 24–34.
Inger, R.F. (1954) Systematics and zoogeography of Philippine Amphibia. Fieldiana: Zoology, 33 (4), 183–531.
https://doi.org/10.5962/bhl.title.5571
International Commission on Zoological Nomenclature [“ICZN”] (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, XXIX + 306 pp.
Iskandar, D.T. (1998) The Amphibians of Java and Bali. Research and Development Centre for Biology–LIPI and GEF Biodiversity Collections Project, Bogor, 117 pp.
IUCN Standards and Petitions Subcommittee (2017) Guidelines for using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. Available from: http://www.iucnredlist.org/documents/RedListGuidelines.pdf (accessed 29 December 2017)
Jerdon, T.C. (1870) Notes on Indian herpetology. part 2. Proceedings of the Asiatic Society of Bengal, March 1870, 66–85.
Jiang, J.P., Yuan, F.R., Xie, F. & Zheng, Z.H. (2003) Phylogenetic relationships of some species and genera in megophryids inferred from partial sequences of mitochondrial 12S and 16S rRNA genes. Zoological Research, 24 (4), 241–248.
Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9 (4), 286-–298.
https://doi.org/10.1093/bib/bbn013Khan, M.M.H. (2008) Protected Areas of Bangladesh; A Guide to Wildlife. Nishorgo Programme, Dhaka, 305 pp.
Kuhl, H. & van Hasselt, J.C. (1822a) Uittreksels uit breieven van de Heeren Kuhl en Van Hasselt, aan de Heeren C. J. Temminck, Th. van Swinderen en W. de Haan. Algemeene Konst-en Letter-Bode, 7, 99–104.
Kuhl, H. & van Hasselt, J.C. (1822b) Aus einem Schreiben von Dr. Kuhl und Dr. Van Hasselt aus Java, an Professor Th. van Swinberen zu Gronigen. Isis von Oken, 10, 472–476.
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/msw054Lau, M.W.N. & Chan, B. (2004) Megophrys mangshanensis. The IUCN Red List of Threatened Species, 2004, e.T57644A11668918. Available from: https://doi.org/10.2305/IUCN.UK.2004.RLTS.T57644A11668918.en (accessed 20 April 2018)
Leaché, A.D. & Fujita, M.K. (2010) Bayesian species delimitation in West African forest geckos (Hemidactylus fasciatus). Proceedings of the Royal Society of London B: Biological Sciences, 277, 3071–3077.
https://doi.org/10.1098/rspb.2010.0662Li, P.-P., Zhao, E.-M. & Dong, B.-J. (2010) Amphibians and Reptiles of Tibet. Science Press, Beijing, XII + 251 pp.
Li, Y.L., Jin, M.J., Zhao, J., Liu, Z., Wang, Y.Y. & Pang, H. (2014) Description of two new species of the genus Megophrys (Amphibia: Anura: Megophryidae) from Heishiding Nature Reserve, Fengkai, Guangdong, China, based on molecular and morphological data. Zootaxa, 3795 (4), 449–471.
https://doi.org/10.11646/zootaxa.3795.4.5Liu, C.-C. (1950) Amphibians of western China. Fieldiana, 2, 1–400, pls. 1–10.
Mahony, S. (2008) Redescription and generic reallocation of Rana mawphlangensis Pillai & Chanda, 1977 (Amphibia: Ranidae). Hamadryad, 33 (1), 1–12.
Mahony, S. (2011) Two new species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae), from western Thailand and southern Cambodia. Zootaxa, 2734, 23–39.
Mahony, S., Sengupta, S., Kamei, R.G. & Biju, S.D. (2011) A new low altitude species of Megophrys Kuhl and Van Hasselt (Amphibia: Megophryidae), from Assam, Northeast India. Zootaxa, 3059, 36–46.
Mahony, S., Teeling, E.C. & Biju, S.D. (2013) Three new species of horned frogs, Megophrys (Amphibia: Megophryidae), from Northeast India, with a resolution to the identity of Megophrys boettgeri populations reported from the region. Zootaxa, 3722 (2), 143–169.
https://doi.org/10.11646/zootaxa.3722.2.2Mahony, S., Foley, N.C., 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 + S.I. 33 pp.
https://doi.org/10.1093/molbev/msw267Mathew, R. & Sen, N. (2007) Description of two new species of Xenophrys (Amphibia: Anura: Megophryidae) from north-east India. Cobra, 1 (2), 18–28.
Mathew, R. & Sen, N. (2010) Pictorial Guide to Amphibians of North East India. Zoological Survey of India, Kolkata, 144 pp.
McLeod, D.S. (2010) Of Least Concern? Systematics of a cryptic species complex: Limnonectes kuhlii (Amphibia: Anura: Dicroglossidae). Molecular Phylogenetics and Evolution, 56, 991–1000.
https://doi.org/10.1016/j.ympev.2010.04.004MCZbase: The Database of the Zoological Collections. Museum of Comparative Zoology, Harvard University (2018) Megophrys major. Available from: https://mczbase.mcz.harvard.edu/guid/MCZ:Herp:A-6814# (accessed 20 January 2018)
Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Institute of Electrical and Electronics Engineers (Eds.), Proceedings of the Gateway Computing Environments Workshop (GCE), Piscataway, New Orleans, pp. 1–8.
https://doi.org/10.1109/GCE.2010.5676129Murphy, W.J. & O'Brien, S.J. (2007) Designing and optimizing comparative anchor primers for comparative gene mapping and phylogenetic inference. Nature Protocols, 2 (11), 3022–3030.
https://doi.org/10.1038/nprot.2007.429Nabhitabhata J., Chan-ard, T. & Chuaynkern, Y. (2000) Checklist of Amphibians and Reptiles in Thailand. Office of Environmental Policy and Planning, Bangkok, 152 pp.
Nanhoe, L.M.R. & Ouboter, P.E. (1987) The distribution of reptiles and amphibians in the Annapurna–Dhaulagiri region (Nepal). Zoologische Verhandelingen, 240, 3–100.
Nylander, J.A.A., Wilgenbusch, J.C., Warren, D.L. & Swofford, D.L. (2008) AWTY (Are We There Yet?): a system for graphical exploration of MCMC convergence in Bayesian phylogenetic inference. Bioinformatics, 24 (4), 581–583.
https://doi.org/10.1093/bioinformatics/btm388Ohler, A., Marquis, O., Swan, S. & Grosjean, S. (2000) Amphibian diversity of Hoang Lien Nature Reserve (Lao Cai Province, northern Vietnam) with description of two new species. Herpetozoa, 13 (1–2), 71–87.
Ohler, A., Swan, S.R. & Daltry, J.C. (2002) A recent survey of the amphibian fauna of the Cardamom Mountains, southwest Cambodia with descriptions of three new species. The Raffles Bulletin of Zoology, 50 (2), 465–481.
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, 1–83.
Orlov, N.L., Murphy, R.W., Ananjeva, N.B., Ryabov, S.A. & Ho, T.C. (2002) Herpetofauna of Vietnam, a checklist. Part 1. Amphibia. Russian Journal of Herpetology, 9 (2), 81–104.
Orlov, N.L., Poyarkov Junior, N.A. & Nguyen, T.T. (2015) Taxonomic notes on Megophrys frogs (Megophryidae: Anura) of Vietnam, with description of a new species. Russian Journal of Herpetology, 22 (3), 206–218.
Palumbi, S.R. (1996) Nucleic acids II: the polymerase chain reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics. 2nd Edition. Sinauer Associates, Sunderland, pp. 205–247.
Pawar, S. & Birand, A. (2001) A Survey of Amphibians, Reptiles, and Birds of Northeast India. CERC Technical Report #6. Center for Ecological Research and Conservation, Mysore, 75 pp.
Pillai, R.S. & Chanda, S.K. (1976) The distribution pattern of Amphibia in Northeast India. Journal of the Assam Science Society, 19 (53), 1–4.
Pillai, R.S. & Chanda, S.K. (1979) Amphibian fauna of Khasi Hills, Meghalaya. Records of the Zoological Survey of India, 75, 383–395.
Pillai, R.S. & Chanda, S.K. (1981) Amphibian fauna of Garo Hills, Meghalaya with description of a new species of Rana. Records of the Zoological Survey of India, 79, 159–168.
Pope, C.H. & Boring, A.M. (1940) A survey of Chinese Amphibia. Peking Natural History Bulletin, 15, 13–86.
Poyarkov Junior, N.A., Rowley, J.J., 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.3931.2.3Poyarkov Junior, N.A., Doung, T.V., Orlov, N.L., Gogoleva, S.S., Vassilieva, A.B., Nguyen, L.T., Nguyen, V.D.H., Nguyen, S.N., Che, J. & Mahony, S. (2017) Molecular, morphological and acoustic assessment of the genus Ophryophryne (Anura, Megophryidae) from Langbian Plateau, southern Vietnam, with description of a new species. Zookeys, 672, 49–120.
https://doi.org/10.3897/zookeys.672.10624Purakayastha J. & Matsui M. (2010) A new species of Fejervarya (Anura: Dicroglossidae) from Mawphlang, northeastern India. Asian Herpetological Research, 3 (1), 31–37.
Pyron, R.A. (2014) Biogeographic analysis reveals ancient continental vicariance and recent oceanic dispersal in amphibians. Systematic Biology, 63 (5), 779–797.
https://doi.org/10.1093/sysbio/syu042Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised 3 classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–83.
https://doi.org/10.1016/j.ympev.2011.06.012Rai, K.R. & Anders, C. (2002) Megophrys robusta (Boulenger, 1908). In: Schleich, H.H. & Kästle, W. (Eds.), Amphibians and Reptiles of Nepal. A.R.G. Gantner Verlag K.G., Ruggel, pp. 167–171.
Rambaut, A. (2009) FigTree version 1.3.1. Computer program distributed by the author. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed January 2012)
Rambaut, A. & Drummond, A.J. (2013) Tracer, version 1.6, MCMC trace analysis package. Available from: http://tree.bio.ed.ac.uk/software/tracer/ (accessed January 2012)
Rannala, B. & Yang, Z. (2003) Bayes estimation of species divergence times and ancestral population sizes using DNA sequences from multiple loci. Genetics, 164, 1645–1656.
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, 539–542.
https://doi.org/10.1093/sysbio/sys029Salerno, P.E., Señaris, J.C., Rojas-Runjaic, F.J.M. & Cannatella, D.C. (2015) Recent evolutionary history of lost world endemics: Population genetics, species delimitation, and phylogeography of sky-island treefrogs. Molecular Phylogenetics and Evolution, 82, 314–323.
https://doi.org/10.1016/j.ympev.2014.10.020Sangma, M.A. & Saikia, P.K. (2015) Checklist of amphibians inventoried from Tura Peak of West Garo Hills, Meghalaya, India. Journal of Global Biosciences, 4 (5), 2404–2424.
Sarkar, A.K. & Ray, S. (2006) Amphibia. In: Alfred, J.R.B. (Ed.), Fauna of Arunachal Pradesh. State Fauna Series. 13 (1). Zoological Survey of India, Kolkata, pp. 285–316.
Sarkar A.K., Biswas, M.L. & Ray, S. (1992) Amphibia. In: “The Director of the Zoological Survey of India” (Ed.), Fauna of West Bengal. State Fauna Series. 3 (2). Zoological Survey of India, Kolkata, pp. 67–100.
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 (2), 111–131.
Schlagintweit, H., Schlagintweit, A. & Schlagintweit, R. (1861) Results of a Scientific Mission to India and High Asia, Undertaken Between the Years MDCCCLIV and MDCCCLVIIL, by Order of the Court of Directors of the Honourable East India Company: With an Atlas of Panoramas, Views, and Maps. Vol. I. Astronomical Determinations of Latitudes and Longitudes and Magnetic Observations During a Scientific Mission to India and High Asia. F.A. Brockhaus, Leipzig and Trübner & Co., London, XII + 494 pp.
Sclater, W.L. (1892a) On some specimens of frogs in the Indian Museum, Calcutta with description of several new species. Proceedings of the Zoological Society of London, 1892, 341–348, pl. XXIV.
Sclater, W.L. (1892b) List of the Batrachia in the Indian Museum. Taylor and Francis, London, VIII + 43 pp.
Sen, N. (2004) Further notes on statewise distribution of the amphibian fauna of Northeast India. Records of the Zoological Survey of India Occasional Papers, 102 (3–4), 105–112.
Sen, N. & Mathew, R. (2008) Bibliographical notes on the amphibians of Northeast India. Records of the Zoological Survey of India Occasional Papers, 291, 1–58.
Setiadi, M.I., McGuire, J.A., Brown, R.M., Zubairi, M., Iskandar, D.T., Andayani, N., Supriatna, J. & Evans, B.J. (2011) Adaptive radiation and ecological opportunity in Sulawesi and Philippine fanged frog (Limnonectes) communities. The American Naturalist, 178 (2), 221-–240.
https://doi.org/10.1086/660830Shen, X.X., Liang, D., Wen, J.Z. & Zhang, P. (2011) Multiple genome alignments facilitate development of NPCL markers: a case study of tetrapod phylogeny focusing on the position of turtles. Molecular Biology and Evolution, 28, 3237–3252.
https://doi.org/10.1093/molbev/msr148Smith, M.A. (1926) The function of the “funnel” mouth of the tadpoles of Megalophrys, with a note on M. aceras Boulenger. Proceedings of the Zoological Society of London, 4, 983–988.
https://doi.org/10.1111/j.1096-3642.1926.tb02229.xSmith, M.A. (1929) On a collection of amphibians and reptiles from the upper reaches of the Brahmaputra. Records of the Indian Museum, 31, 77–80.
Smith, M.A. (1940) The amphibians and reptiles obtained by Mr. Ronald Kaulback in Upper Burma. Records of the Indian Museum, 42, 465–486, pl. VIII.
Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033Stephens, M. & Scheet, P. (2005) Accounting for decay of linkage disequilibrium in haplotype inference and missing-data imputation. The American Journal of Human Genetics, 76 (3), 449– 462.
https://doi.org/10.1086/428594Stephens, M., Smith, N. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. The American Journal of Human Genetics, 68 (4), 978–989.
https://doi.org/10.1086/319501Stuart, B.L. (1999) Amphibians and reptiles. In: Duckworth, J.W., Salter, R.E. & Khounboline, K. (Eds.), Wildlife in Lao PDR. 1999 Status Report. IUCN-The World Conservation Union/Wildlife Conservation Society/Centre for Protected Areas and Watershed Management, Vientiane, pp. 43–67.
Stuart, B.L. (2005) New frog records from Laos. Herpetological Review, 36 (4), 473–479.
Stuart, B.L., Chuaynkern, Y., Chan-ard, T. & Inger, R.F. (2006) Three new species of frogs and a new tadpole from eastern Thailand. Fieldiana, 111, 1–19.
https://doi.org/10.3158/0015-0754(2006)187[1:TNSOFA]2.0.CO;2Stuart, B.L., Rowley, J.J.L., Neang, T., Emmett, D.A. & Sitha, S. (2010) Significant new records of amphibians and reptiles from Virachey National Park, northeastern Cambodia. Cambodian Journal of Natural History, 2010 (1), 38–47.
Subba, B., Aravind, N.A. & Ravikanth, G. (2016) Amphibians of the Sikkim Himalaya, India: an annotated checklist. Check List, 13 (1), 1–14.
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
https://doi.org/10.1093/molbev/mst197Tapley, B., Cutajar, T., Mahony, S., Nguyen, C.T., Dau, V.Q., Nguyen, T.T., Luong, H.V. & Rowley, J.J.L. (2017) The Vietnamese population of Megophrys kuatunensis (Amphibia: Megophryidae) represents a new species of Asian horned frog from Vietnam and southern China. Zootaxa, 4344 (3), 465–492.
https://doi.org/10.11646/zootaxa.4344.3.3Tapley, B., Cutajar, T., Mahony, S., Nguyen, C.T, Dau, V.Q., Luong, A.M., Le, D.T., Nguyen, T.T., Nguyen, T.Q., Portway, C., Luong, H.V. & Rowley, J.J.L. (2018) Two new and potentially highly threatened Megophrys Horned frogs (Amphibia: Megophryidae) from Indochina’s highest mountains. Zootaxa, 4508 (3), 301–333.
https://doi.org/10.11646/zootaxa.4508.3.1
Taylor, E.H. (1962) The amphibian fauna of Thailand. The University of Kansas Science Bulletin, 43, 255–599.
https://doi.org/10.5962/bhl.part.13347Tron, R.K.L., Das, I., Hooroo, R.N.K. & Rangad, D. (2015) Spring-breeding and reproductive mode in Leptolalax khasiorum (Anura, Megophryidae) in northeastern India. Russian Journal of Herpetology, 22 (3), 224–232.
Wagler, J.G. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Säugethiere und Vögel. J.G. Cotta, München, VI + 134 pp.
Wang, Y.Y., Zhang, T.D., Zhao, J., Sung, Y.H., Yang, J.H., Pang, H. & Zhang, Z. (2012) Description of a new species of the genus Xenophrys Günther, 1864 (Amphibia: Anura: Megophryidae) from Mount Jinggang, China, based on molecular and morphological data. Zootaxa, 3546, 53–67.
Wang, Y., Zhao, J., Yang, J., Zhou, Z., Chen, G. & Liu, Y. (2014) Morphology, molecular genetics, and bioacoustics support two new sympatric Xenophrys toads (Amphibia: Anura: Megophryidae) in Southeast China. PLOS ONE, 9 (4), e93075.
https://doi.org/10.1371/journal.pone.0093075Wangyal, J.T. (2013) New records of reptiles and amphibians from Bhutan. Journal of Threatened Taxa, 5, 4774–4783.
https://doi.org/10.11609/JoTT.o3539.4774-83Wangyal, J.T. & Gurung, D.B. (2012) Amphibians of Punakha-Wangdue Phodrang Valley, Bhutan. Frog Leg Newsletter of the Amphibian Network of South Asia and Amphibian Specialist Group-South Asia, 18, 31–44.
Wogan, G.O.U., Vindum, J.V., Wilkinson, J.A., Koo, M.S., Slowinski, J.B., Win, H. Thin, T. Kyi, S.W., Oo, S.L., Lwin, K.S. & Shein, A.K. (2008) New country records and range extensions for Myanmar amphibians and reptiles. Hamadryad, 33, 83–96.
Yang, D.-T., Liu, W.-Z., Rao, D.-Q. & He, Y.-H. (1994) Studies on the diversity of three populations of Megophrys lateralis. Zoological Research, 15, 124–130.
Yang, J.-h., Wang, J. & Wang, Y.-y. (2018) A new species of the genus Megophrys (Anura: Megophryidae) from Yunnan Province, China. Zootaxa, 4413, 325–338.
https://doi.org/10.11646/zootaxa.4413.2.5
Yang, Z. & Rannala, B. (2010) Bayesian species delimitation using multilocus sequence data. Proceedings of the National Academy of Sciences, 107 (20), 9264–9269.
https://doi.org/10.1073/pnas.0913022107Ye, C.-Y. & Fei, L. (1992) A new pelobatid toad of the genus Megophrys from Xizang, China. Acta Herpetologica Sinica, 1–2, 50–52.
Zhang, P., Papenfuss, T.J., Wake, M.H., Qu, L. & Wake, D.B. (2008) Phylogeny and biogeography of the family Salamandridae (Amphibia: Caudata) inferred from complete mitochondrial genomes. Molecular Phylogenetics and Evolution, 49, 586–597.
https://doi.org/10.1016/j.ympev.2008.08.020Zhang, Y., Li, G., Xiao, N., Li, J., Pan, T, Wang, H., Zhang, B. & Zhou, J. (2017) A new species of the genus Xenophrys (Amphibia: Anura: Megophryidae) from Libo County, Guizhou, China. Asiatic Herpetological Research, 8, 75–85.
Zhao, E.-m. & Adler, K. (1993) Herpetology of China. SSAR, Oxford, 552 + V pp., pls. 1–48 + 1.
Zhao, E. & Li, S. (1987) The herpetofaunal composition and characteristics of Mount Namjagbarwa, Tibet. Acta Herpetologica Sinica, 6 (2), 36–42.
Zhao, W, Rao, D., Lü, S. & Dong, B. (2005) Herpetological surveys of Xizang Autonomous Region. 2. Medog. Sichuan Journal of Zoology, 24 (3), 250–253.
Zheng, Y.-C., Mo, B.-H., Liu, Z.-J. & Xeng, X.-M. (2004a) Phylogenetic relationships of megophryid genera (Anura: Megophryidae) based on partial sequences of mitochondrial 16 rRNA gene. Zoological Research, 25 (3), 205–213.
Zheng, Y.-C., Xeng, X.-M., Yuan, Y.-Z. & Liu, Z.-J. (2004b) Phylogenetic position of Ophryophryne and four Leptobrachium group genera in Megophryidae (Anura). Sichuan Journal of Zoology, 23 (3), 290–295.