Abstract
We evaluated the taxonomic status of the genus Megophrys in Sumatra using molecular and morphological data. Mitochondrial phylogenetic inference and morphological data revealed two undescribed species, one in southern Sumatra—M. selatanensis sp. nov. and one in northern Sumatra—M. acehensis sp. nov. We also detected a potential cryptic species within M. parallela, but refrain from describing this lineage here due to insufficient data. Genetic variation within Sumatran Megophrys is highly structured and will require additional geographic sampling to understand the interplay between geography and genetics in Sumatran Megophrys.
References
Arifin, U., Chan, K.O., Smart, U., Hertwig, S.T., Smith, E.N., Iskandar, D.T. & Haas, A. (2021) Revisiting the phylogenetic predicament of the genus Huia (Amphibia: Ranidae) using molecular data and tadpole morphology. Zoological Journal of the Linnean Society, 193 (2), 673–699, https://doi.org/10.1093/zoolinnean/zlaa158
Barley, A.J., White, J., Diesmos, A.C. & Brown, R.M. (2013) The challenge of species delimitation at the extremes: diversification without morphological change in Philippine sun skinks. Evolution, 67 (12), 3556–3572. https://doi.org/10.1111/evo.12219
Boulenger, G.A. (1885) Description of a new frog of the genus Megalophrys. Proceedings of the scientific meeting of the Zoological Society of London, 53 (4), 850–850. https://doi.org/10.1111/j.1096-3642.1885.tb02927.x
Boulenger, G.A. (1899) Description of three new reptiles and a new batrachian from Mount Kinabalu, North Borneo. Annals and Magazine of Natural History, 7 (4), 451−454. https://doi.org/10.1080/00222939908678228
Boulenger, G.A. (1903) Report on the batrachians and reptiles. In: Annandale, N. & Robinson, H.C. (Eds.), Fasciculi Malayenses. Zoology. Part I. Anthropological and Zoological Results of an Expedition to Perak and the Siamese Malay States 1901–1902. Longmans, Green & Co, London, pp. 131–176.
Brown, R.M., Siller, C.C., Diesmos, A.C. & Alcala, A.C. (2009) Philippine frogs of the genus Leptobrachium (Anura: Megophrydae): phylogeny-based species delimitation, taxonomic review, and description of three new species. Herpetological Monographs, 23 (1), 1–44. https://doi.org/10.1655/09-037.1
Chan, K.O., Hutter, C.R., Wood, P.L., Grismer, L.L., Das, I. & Brown, R.M. (2020) Gene flow creates a mirage of cryptic species in a Southeast Asian spotted stream frog complex. Molecular Ecology, 29 (20), 3970–3987. https://doi.org/10.1111/mec.15603
Chan, K.O., Hutter, C.R., Wood, P.L.J., Su, Y.C. & Brown, R.M. (2021) Gene Flow Increases Phylogenetic Structure and Inflates Cryptic Species Estimations: A Case Study on Widespread Philippine Puddle Frogs (Occidozyga laevis). Systematic Biology, syab034. [published online] https://doi.org/10.1093/sysbio/syab034
De Bruyn, M., Stelbrink, B., Morley, R.J., Hall, R., Carvalho, G.R., Cannon, C.H., van den Bergh, G., Meijaard, E., Metcalfe, I. & Boitani, L. (2014) Borneo and Indochina are major evolutionary hotspots for Southeast Asian biodiversity. Systematic Biology, 63 (6), 879–901. https://doi.org/10.1093/sysbio/syu047
Chen, J.M., Zhou, W.W., Poyarkov, N.A., Stuart, B.L., Brown, R.F., Lathrop, A., Wang, Y.Y., Yuan, Z.Y., Jiang, K., Hou, M., Chen, H.M., Suwannapoom, C., Nguyen S.N., Duong, T.V., 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.2016.09.004
Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24 (1−4), 6−21.
Diesmos, A.C., Watters, J.L., Huron, N.A., Davis, D.R., Alcala, A.C., Crombie R.I., Afuang, L.E., Gee-das, G., Sison, R.V., Sanguila, M.B., Penrod, M.L., Labonte, M.J., Davey C.S., Leone E.A., Diesmos M.L., Sy, M.Y., Welton, L.J., Brown R.M. & Siler C.D. (2015) Amphibians of the Philippines, Part I: Checklist of the Species. Proceedings of the California Academy of Science, 62 (20), 457−539.
Dubois, A. (1987) Miscellanea taxonomica Batrachologica (I). Alytes, 5, 7–95.
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.
Dubois, A., Ohler, A. & Pyron, R.A. (2021) New concepts and methods for phylogenetic taxonomy and nomenclature in zoology, exemplified by a new ranked cladonomy of recent amphibians (Lissamphibia). Megataxa, 5 (1), 1–738. https://doi.org/10.11646/megataxa.5.1.1
Eto, K., Matsui, M. & Nishikawa, K. (2015) Description of a new species of the genus Leptobrachella (Amphibia, Anura, Megophryidae) from Borneo. Current Herpetology, 34 (2), 128–139. https://doi.org/10.5358/hsj.34.128
Fei, L. & Ye, C.Y. (2016) Amphibians of China. Vol. 1. Science Press, Beijing, 1040 pp.
Foquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmel, N.J. (2007) Underestimation of species richness in Neotropical frog revealed by mtDNA analyses. PLoSONE, 2 (10), 1–10. https://doi.org/10.1371/journal.pone.0001109
Frost, D.R. (2021) Amphibian Species of the World, an Online Reference. Version 6.0. Electronic Database, American Museum of Natural History, New York. Available from, http,//research.amnh.org/herpetology/amphibia/ index.html (accessed 8 July 2021)
Günther, A.C.L.G. (1864) The reptiles of British India. The Ray Society, London, 452 pp.
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.
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/10.14203/treubia.v44i0.3286
Hamidy, A., Munir, M., Mumpuni, M., Rahmania, M. & Kholik, A.A. (2018) Detection of cryptic taxa in the genus Leptophryne (Fitzinger, 1843) (Amphibia: Bufonidae) and the description of a new species from Java, Indonesia. Zootaxa, 4450 (4), 427–444. https://doi.org/10.11646/zootaxa.4450.4.2
Hasan, M., Islam, M.M., Khan, Md. M.M., Igawa, T., Alam, M.S., Djong, H.T., Kurniawan, N., Joshy, H., Sen, Y.H., Belabut, D.M., Kurabayashi, A., Kuramoto, M. & Sumida, M. (2014) Genetic divergence of South and Southeast Asian frogs; a case study of several taxa based on 16S RNA gene data with notes on the generic name Fejervarya. Turkish Journal of Zoology, 38 (4), 389–411. https://doi.org/10.3906/zoo-1308-36
Harvey, M.B., Hamidy, A., Kurniawan, N., Shaney, K. & Smith, E.N. (2014) Three new species of Pseudocalotes (Squamata: Agamidae) from southern Sumatra, Indonesia. Zootaxa, 3841 (2), 211–238. https://doi.org/10.11646/zootaxa.3841.2.3
Harvey, M.B., O’Connell, K.A., Wostl, E., Riyanto, A., Kurniawan, N., Smith, E.N. & Grismer, L.L. (2016) Redescription Cyrtodactylus lateralis (Werner) (Squamata: Gekkonidae) and phylogeny of the prehensile-tailed Cyrtodactylus. Zootaxa, 4107 (4), 517–540. https://doi.org/10.11646/zootaxa.4107.4.3
Harvey, M.B., Shaney, K., Sidik, I., Kurniawan, N. & Smith, E.N. (2017a) Endemic dragons of Sumatra’s volcanoes: New species of Dendragama (Squamata: Agamidae) and status of Salea rosaceum Thominot. Herpetological Monographs, 31 (1), 69–97. https://doi.org/10.1655/HERPMONOGRAPHS-D-16-00012
Harvey, M.B., Shaney, K., Hamidy, A., Kurniawan, N. & Smith, E.N. (2017b) A new species of Pseudocalotes (Squamata: Agamidae) from the Bukit Barisan Range of Sumatra with an estimation of its phylogeny. Zootaxa, 4276 (2), 215–232. https://doi.org/10.11646/zootaxa.4276.2.4
Hedges, S.B., Nussbaum, R.A. & Maxson, L.R. (1993) Caecilian phylogeny and biogeography inferred from mitochondrial DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia: Gymnophiona). Herpetologica Monograph, 7, 64–76.
https://doi.org/10.2307/1466952
Hu, C.C., Wu, Y.Q., Ma, L., Chen, Y.J. & Ji, X. (2019) Genetic and morphological divergence among three closely related Phrynocephalus species (Agamidae). BMC Ecology and Evolution, 19, 114. https://doi.org/10.1186/s12862-019-1443-y
Inger, R.F. (1954) The systematics and zoogeography of Philippine Amphibia. Fieldiana Zoology, 33 (4), 183–531. https://doi.org/10.5962/bhl.title.5571
Inger, R.F. (1966) The systematic and zoogeography of the amphibians of Borneo. Fieldiana Zoology, 52, 1–402. https://doi.org/10.5962/bhl.title.3147
Inger, R.F. (1989) Four new species of frogs from Borneo. Malayan Nature Journal, 42, 229–243.
Inger, R.F., Stuebing, R.B. & Tan, F.L. (1995) New species and new records of anurans from Borneo. The Raffles Bulletin of Zoology, 43 (1), 115−131.
Inger, R.F. & Iskandar, D.T. (2005) A collection of amphibians from West Sumatra, with description of new species of Megophrys (Amphibia: Anura). The Raffles Bulletin of Zoology, 53 (1), 133–142.
Inger, R.F. & Stuebing, R.B. (2005) A field guide to the frogs of Borneo. 2nd Edition. Natural History Publication Borneo Sdn. Bhd. Kota Kinabalu, Sabah, 201 pp.
Inger, R.F., Stuebing, R.B., Grafe, T.U. & Dehling, J.M. (2017). A field guide to the Frogs of Borneo. Third edition. Natural History Publication Borneo. Kinabalu, 236 pp.
Katoh, K. & Standley, D.M. (2013) MAFFT Multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 772–780. https://doi.org/10.1093/molbev/mst010
Kurniawan, N., Affandi, A., Wardani, E.E., Chomsy, I.N., Firdaus, A.S. & Kurnianto, A.S. (2017) Phylogenetic patterns of genus Megophrys (Anura: Megophryidae) from Java and Sumatra. Journal of Experimental Life Science, 7, 76–83. https://doi.org/10.21776/ub.jels.2017.007.02.03
Lathrop, A. (1997) Taxonomic review of the Megophryid frogs (Anura: Pelobatidae). Asiatic Herpetological Research, 7, 68–79. https://doi.org/10.5962/bhl.part.18857
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 (1), 44–68. https://doi.org/10.1080/106351502753475871
Li, Y., Zhang, D.D., Lyu, Z.T., Wang, J., Li, Y.L., Liu, Z.Y., Chen, H.H., Rao, D.D., Jin, Z.F., Zhang, C.Y. & Wang Y.Y. (2020) Review of the genus Brachytarsophrys (Anura: Megophryidae), with revalidation of Brachytarsophrys platyparietus and description of a new species from China. Zoological Research, 41 (2), 105–122. https://doi.org/10.24272/j.issn.2095-8137.2020.033
Liu, Z., Chen, G., Zhu, T., Zeng, Z., Lyu, Z., Wang, J., Messenger, K., Greenberg, A.J., Guo, Z., Yang, Z., Shi, S. & Wang, Y. (2018) Prevalence of cryptic species in morphologically uniform taxa—Fast speciation and evolutionary radiation in Asian frogs. Molecular Phylogenetics and Evolution, 127, 723−731. https://doi.org/10.1016/j.ympev.2018.06.020
Lohman, D.J., de Bruyn, M., Page, T., von Rintelen, K., Hall, R., Ng, P.K. & von Rintelen, T. (2011) Biogeography of the Indo-Australian archipelago. Annual Review of Ecology, Evolution, and Systematics, 42, 205–226. https://doi.org/10.1146/annurev-ecolsys-102710-145001
Luo, T., Wang, Y., Wang, S., Lu, X., Wang, W., Deng, H. & Zhou, J. (2021) A species of the genus Panophrys (Anura, Megophryidae) from southeastern Guizhou Province, China. ZooKeys, 1047, 27–60. https://doi.org/10.3897/zookeys.1047.61097
Lyu, Z.T., Li, Y.Q., Zeng, S.C., Zhao, J., Liu, Z.Y., Guo, G.X. & Wang, Y.Y. (2020) Four new species of Asian horned toads (Anura, Megophryidae, Megophrys) from southern China. ZooKeys, 942, 105–140. https://doi.org/10.3897/zookeys.942.47983
Lyu, Z.T., Zeng, Z., Wang, J., Liu, Z.Y., Huang, Y.Q., Li, W.Z. & Wang, Y.Y. (2021) Four new species of Panophrys (Anura, Megophryidae) from eastern China, with discussion on the recognition of Panophrys as a distinct genus. Zootaxa, 4927 (1), 9–40. https://doi.org/10.11646/zootaxa.4927.1.2
Mahony, S., Foley, N.M., Biju, S.D. & Teeling, E.C. (2017) Evolutionary history of the Asian horned frogs (Megophryinae), Integrative approaches to time tree dating in the absence of fossil record. Molecular Biology and Evolution, 34 (3), 744–771. https://doi.org/10.1093/molbev/msw267
Mahony, S., Kamei, R.G., Teeling, E.C. & Biju, S.D. (2018) Cryptic diversity within the Megophrys major species group (Amphibia: Megophryidae) of the Asian Horned Frogs: Phylogenetic perspectives and a taxonomic revision of South Asian taxa, with descriptions of four new species. Zootaxa, 4523 (1), 1–96. https://doi.org/10.11646/zootaxa.4523.1.1
Mahony, S., Kamei, R.G., Teeling, E.C. & Biju, S.D. (2020) Taxonomic review of the Asian Horned Frogs (Amphibia: Megophrys Kuhl & Van Hasselt) of Northeast India and Bangladesh previously misidentified as M. parva (Boulenger), with descriptions of three new species. Journal of Natural History, 54 (1–4), 119–194. https://doi.org/10.1080/00222933.2020.1736679
Malkmus, R. & Matsui, M. (1997) Megophrys kobayashii, ein neuer pelobatider frosch vom Mount Kinabalu, Sauria, 19, 31–37.
Minh, B.Q., Nguyen, M.A.T. & Haeseler, A. (2013) Ultrafast Approximation for phylogenetic bootstrap. Molecular Biology and Evolution, 30 (5), 1188–1195. https://doi.org/10.1093/molbev/mst024
Munir, M., Hamidy, A., Farajallah, A. & Smith, E.N. (2018) A new Megophrys Kuhl and van Hasselt (Amphibia: Megophryidae) from southwestern Sumatra, Indonesia. Zootaxa, 4442 (3), 389−412. https://doi.org/10.11646/zootaxa.4442.3.3
Munir, M., Hamidy, A., Matsui, M., Iskandar, D.T., Sidik, I. & Shimada, T. (2019) A new species of Megophrys Kuhl & van Hasselt (Amphibia: Megophryidae) from Borneo allied to M. nasuta (Schlegel, 1858). Zootaxa, 4679 (1), 1−24. https://doi.org/10.11646/zootaxa.4679.1.1
Nguyen, T.Q., Pham, C.T., Nguyen, T.T., Luong, A.M. & Ziegler, T. (2020). A new species of Megophrys (Amphibia: Anura: Megophryidae) from Vietnam. Zootaxa, 4722 (5), 401–422. https://doi.org/10.11646/zootaxa.4722.5.1
Nosil P. (2012) Ecological speciation. Oxford University Press, Oxford, 280 pp. https://doi.org/10.1093/acprof:osobl/9780199587100.001.0001
Novia, Sanjaya, M.& Gusman, D. (2015) Distribusi Vertikal Anura di Gunung Seblat, Kabupaten Lebong, Bengkulu. In: Prosiding Semirata 2015 Bidang MIPA BKS-PTN Barat. Universitas Tanjungpura, Pontianak, pp. 173–178. [in Indonesian]
O’Connell, K.A., Smith, E.N., Shaney, K.A., Arifin, U., Kurniawan, N., Sidik, I. & Fujita, M.K. (2017) Coalescent species delimitation of a Sumatran parachuting frog. Zoological Scripta, 47 (1), 33–43. https://doi.org/10.1111/zsc.12248
O’Connell, K.A., Hamidy, A., Kurniawan, N.A., Smith, E.N. & Fujita, M.K. (2018a) Synchronous diversification of parachuting frogs (Genus Rhacophorus) on Sumatra and Java. Molecular Phylogenetics and Evolution, 123, 101−112. https://doi.org/10.1016/j.ympev.2018.02.003
O'Connell, K.A., Smart, U., Smith, E.N., Hamidy, A., Kurniawan, N. & Fujita, M. K. (2018b) Within-island diversification underlies parachuting frog (Rhacophorus) species accumulation on the Sunda Shelf. Journal of Biogeography, 45 (4), 929−940. https://doi.org/10.1111/jbi.13162
Oberhummer, E., Barten, C., Schweizer, M., Das, I., Haas, A. & Hertwig, S.T. (2014) Description of the tadpole 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
Poyarkov, N.A. Jr., Duong, T.V., Orlov, N.L., Gogoleva, S.I., 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.672.10624
Rao, D.Q. & Yang, D.T. (1997) The karyotypes of Megophryinae (Pelobatidae) with discussion on their classification and phylogenetic relationships. Asiatic Herpetological Research, 7, 93–102. https://doi.org/10.5962/bhl.part.18858
Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mix models. Bioinformatics, 19 (12), 1572–1574. https://doi.org/10.1093/bioinformatics/btg180
Sanguila, M.B., Cobb, K.A, Siller C.D., Diesmos, A.C., Alcala A.C. & Brown R.M. (2016) The amphibians and reptiles of Mindanao Island, southern Philippines, II: the herpetofauna of northeast Mindanao and adjacent islands. ZooKeys, 624, 1−32. https://doi.org/10.3897/zookeys.624.9814
Shaney, K.J., Maldonado, J., Smart, U., Thamachoti, P., Fujita, M., Hamidy, A., Kurniawan, N., Harvey, M.B. & Smith, E.N. (2020) Phylogeography of montane dragons could shed light on the history of forests and diversification processes on Sumatra. Molecular Phylogenetic and Evolution, 149, 106840. https://doi.org/10.1016/j.ympev.2020.106840
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution, 30 (12), 2725–2729. https://doi.org/10.1093/molbev/mst197
Tanabe, A.S. (2011) Kakusan4 and Aminosan: two programs for comparing nonpartitioned, proportional and separate models for combined molecular phylogenetic analyses of multilocus sequence data. Molecular Ecology Resources, 11 (5), 914–921. https://doi.org/10.1111/j.1755-0998.2011.03021.x
Tapley, B., Cutajar, T.P., 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
Tapley, B., Cutajar, T.P., Nguyen, L.T., Portway, C., Mahony, S., Nguyen, C.T., Harding, L., Luong, H.V. & Rowley, J.J.L. (2021) A new potentially Endangered species of Megophrys (Amphibia: Megophryidae) from Mount Ky Quan San, north-west Vietnam. Journal of Natural History, 54 (39–40), 2543–2575. https://doi.org/10.1080/00222933.2020.1856952
Taylor, E.H. (1920) Philippine Amphibia. The Philippine Journal of Science, 16 (3), 213–359. https://doi.org/10.5962/bhl.part.4751
Taylor, E.H. (1962) The amphibian fauna of Thailand. The University of Kansas Science Bulletin, 43 (8), 265–599. https://doi.org/10.5962/bhl.part.13347
Tian, W. & Hu, Q. (1983) Taxonomic study on genus Megophrys with descriptions of two new genera. Acta Herpetologica Sinica, 2 (2), 41–48.
Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research, 44 (W1), W232–W235. https://doi.org/10.1093/nar/gkw256
Vences, M., Thomas, M., van der Meijden, A., Chiari, Y. & Vieites, D.R. (2005) Comparative performance of the 16S rRNA gene in DNA bar-coding of amphibians. Frontiers in Zoology, 2, 5. https://doi.org/10.1186/1742-9994-2-5
Wang, J., Lyu, Z.T., Liu, Z.Y., Liao, C.K., Zeng, Z.C., Zhao, J., Li, Y.L. & Wang, Y.Y. (2019) Description of six new species of the subgenus Panophrys within the genus Megophrys (Anura, Megophryidae) from southeastern China-based on molecular and morphological data. ZooKeys, 851, 113–164. https://doi.org/10.3897/zookeys.851.29107