Abstract
The diversity of Panophrys horned toads is considered highly underestimated with a large number of undescribed cryptic species. In this work, we describe four Panophrys species from eastern China which were proposed as cryptic species by molecular data in previous study, additionally provide new information on the biogeography of these four species. Panophrys daiyunensis sp. nov. from southern Fujian, Panophrys daoji sp. nov. from eastern Zhejiang, Panophrys sanmingensis sp. nov. from the hilly area among Fujian, Jiangxi and Guangdong, and Panophrys tongboensis sp. nov. from northeastern Jiangxi, can be distinguished from all recognized congeners by a combination of morphological characteristics. The descriptions of these four new species take the recognized species of Panophrys to 51, which is the largest genus within the Asian horned toads subfamily Megophryinae. Considered as an appropriate arrangement for the Asian horned toads currently and applied in this study to describe the new species, the generic recognition of Panophrys is also discussed.
References
AmphibiaWeb (2020) AmphibiaWeb. University of California, Berkeley, California. Available from: http://amphibiaweb.org (accessed 19 June 2020)
Arifin, U., Smart, U., Hertwig, S.T., Smith, E.N., Iskandar, D.T. & Haas, A. (2018) Molecular phylogenetic analysis of a taxonomically unstable ranid from Sumatra, Indonesia, reveals a new genus with gastromyzophorous tadpoles and two new species. Zoosystematics and Evolution, 94, 163–193.
https://doi.org/10.3897/zse.94.22120
Avise, J.C., Nelson, W.S. & Sugita, H. (1994) A speciational history of ‘‘living fossils’’: molecular evolutionary history in horseshoe crabs. Evolution, 48, 1986–2001.
https://doi.org/10.1111/j.1558-5646.1994.tb02228.x
Boulenger, G.A. (1903) Descriptions of three new batrachians from Tonkin. Annals and Magazine of Natural History, Series 7, 12, 186–188.
https://doi.org/10.1080/00222930308678835
Chen, J.M., Zhou, W.W., Poyarkov Jr., 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., 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
Cherty, L.M., Case, S.M. & Wilson, A.C. (1978) Frog perspective on the morphological difference between humans and chimpanzees. Science, 200, 209–211.
https://doi.org/10.1126/science.635583
Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24, 6–21.
Deuti, K., Grosjean, S., Nicolas, V., Vasudevan, K. & Ohler, A. (2017) Nomenclatural puzzle in early Xenophrys nomina (Anura, Megophryidae) solved with description of two new species from India (Darjeeling hills and Sikkim). Alytes, 34, 20–48.
Dubois, A. (1987) Miscellanea taxinomica batrachologica (I). Alytes, 5, 7–95.
Duellman, W.E., Marion, A.B. & Hedges, S.B. (2016) Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae). Zootaxa, 4104 (1), 1–109.
https://doi.org/10.11646/zootaxa.4104.1.1
Fei, L., Hu, S.Q., Ye, C.Y. & Huang, Y.Z. (2009) Fauna Sinica. Amphibia (Vol. 2). Anura. Science Press, Beijing. [in Chinese]
Fei, L. & Ye, C.Y. (2016) Amphibians of China. Vol. 1. Science Press, Beijing, 1040 pp.
Frost, D.R. (2020) Amphibian Species of the World: An Online Reference. Version 6.0. American Museum of Natural History, New York. Available from: http://research.amnh.org/herpetology/amphibia/index.html. (accessed 19 June 2020)
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F., De Sa, R.O., 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, 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
Li, C. & Wang, Y.Z. (2008) Taxonomic review of Megophrys and Xenophrys, and a proposal for Chinese species (Megophryidae, Anura). Acta Zootaxonomica Sinica, 33, 104–106. [in Chinese with English abstract]
Li, J.T., Che, J., Bain, R.H., Zhao, E.M. & Zhang, Y.P. (2008) Molecular phylogeny of Rhacophoridae (Anura): A framework of taxonomic reassignment of species within the genera Aquixalus, Chiromantis, Rhacophorus, and Philautus. Molecular Phylogenetics and Evolution, 48, 302–312.
https://doi.org/10.1016/j.ympev.2008.03.023
Li. S., Xu. N., Liu. J., Jiang. J., Wei. G. & Wang, B. (2018) A New Species of the Asian Toad Genus Megophrys sensu lato (Amphibia: Anura: Megophryidae) from Guizhou Province, China. Asian Herpetological Research, 9, 224–239.
https://doi.org/10.16373/j.cnki.ahr.180072
Li, Y., Zhang, D.D., Lyu, Z.T., Wang, J., Li, Y.L., Liu, Z.Y., Chen, H.H., Rao, D.Q., 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, 105–122.
https://doi.org/10.24272/j.issn.2095-8137.2020.033
Li, Y.L., Jin, M.J., Zhao, J., Liu, Z.Y., 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.3795.4.5
Liu, J., Li, S., Wei, G., Xu, N., Cheng, Y., Wang, B. & Wu, J. (2020) A New Species of the Asian Toad Genus Megophrys sensu lato (Anura: Megophryidae) from Guizhou Province, China. Asian Herpetological Research, 11, 1–18.
https://doi.org/10.16373/j.cnki.ahr.190041
Liu, Z.Y., Chen, G.L., Zhu, T.Q., Zeng, Z.C., Lyu, Z.T., Wang, J., Messenger, K., Greenberg, A.J., Guo, Z.X., Yang, Z.H., Shi, S.H. & Wang, Y.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
Lyu, Z.T., Li, Y.Q., Zeng, Z.C., Zhao, J., Liu, Z.L., 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.C., Wang, J., Lin, C.Y., Liu, Z.Y. & Wang, Y.Y. (2017) Resurrection of genus Nidirana (Anura: Ranidae) and synonymizing N. caldwelli with N. adenopleura, with description of a new species from China. Amphibia-Reptilia, 38, 483–502.
https://doi.org/10.1163/15685381-00003130
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, 744–771. https://doi.org/10.1093/molbev/msw267
Mayr, E. (1942) Systematics and the Origin of Species from the Viewpoint of a Zoologist. Columbia University Press, New York, xiv + 334 pp.
Messenger, K.R., Dahn, H.A., Liang, Y., Xie, P., Wang, Y. & Lu, C. (2019) A new species of the genus Megophrys Gunther, 1864 (Amphibia: Anura: Megophryidae) from Mount Wuyi, China. Zootaxa, 4554 (2), 561–583.
https://doi.org/10.11646/zootaxa.4554.2.9
Rao, D.Q. & Yang, D.T. (1997) The karyotypes of Megophryinae (Pelobatinae) with a discussion on their classification and phylogenetic relationships. Asiatic Herpetological Research, 7, 93–102.
https://doi.org/10.5962/bhl.part.18858
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/sys029
Schopf, T.J. (1984). Rates of Evolution and the Notion of” Living Fossils”. Annual Review of Earth and Planetary Sciences, 12, 245–292.
https://doi.org/10.1146/annurev.ea.12.050184.001333
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/mst197
Tapley, 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.4344.3.3
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
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
Vences, M., Guayasamin, J.M., Miralles, A. & De la Riva, I. (2013) To name or not to name: criteria to promote economy of change in supraspecific Linnean classification schemes. Zootaxa, 3636 (2), 201–244.
https://doi.org/10.11646/zootaxa.3636.2.1
Wang, B., Wu, Y.Q., Peng, J.W., Shi, S.C., Lu, N.N. & Wu, J. (2020) A new Megophrys Kuhl & Van Hasselt (Amphibia, Megophryidae) from southeastern China. ZooKeys, 904, 35–62.
https://doi.org/10.3897/zookeys.904.47354
Wang, J., Liu, Z.Y., Lyu, Z.T., Zeng, Z.C. & Wang, Y.Y. (2017a) A new species of the genus Xenophrys (Amphibia: Anura: Megophryidae) from an offshore island in Guangdong Province, southeastern China. Zootaxa, 4324 (3), 541–556.
https://doi.org/10.11646/zootaxa.4324.3.8
Wang, J., Lyu, Z.T., Liu, Z.Y., Liao, C.K., Zeng, Z.C., Zhao, J., Li, Y.L. & Wang, Y.Y. (2019a) 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
Wang, L., Deng, X., Liu, Y., Wu, Q. & Liu, Z. (2019b) A new species of the genus Megophrys (Amphibia: Anura: Megophryidae) from Hunan, China. Zootaxa, 4695 (4), 301–330.
https://doi.org/10.11646/zootaxa.4695.4.1
Wang, Y.F., Liu, B.Q., Jiang, K., Jin, W., Xu, J.N. & Wu, C.H. (2017b) A new species of the Horn Toad of the genus Xenophrys from Zhejiang, China (Amphibia: Megophryidae). Chinese Journal of Zoology, 52, 19–29. [in Chinese with English abstract]
https://doi.org/10.13859/j.cjz.201701003
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 (1), 53–67.
https://doi.org/10.11646/zootaxa.3546.1.4
Wang, Y.Y., Zhao, J., Yang, J.H., Zhou, Z., Chen, G.L. & Liu, Y. (2014) Morphology, molecular genetics, and bioacoustics support two new sympatric Xenophrys toads (Amphibia: Anura: Megophryidae) in Southeast China. PloS One, 9, e93075.
https://doi.org/10.1371/journal.pone.0093075
Wilson, A.C., Carlson, S.S. & White, T.J. (1977) Biochemical evolution. Annual review of biochemistry, 46, 573–639.
https://doi.org/10.1146/annurev.bi.46.070177.003041
Wu, Y.H., Suwannapoom, C., Poyarkov Jr., N.A., Paawangkhanat, P., Xu, K., Jin, J.Q., Murphy, R.W. & Che, J. (2019) A new species of the genus Xenophrys (Anura:Megophryidae) from northern Thailand. Zoological Research, 40, 564–574.
https://doi.org/10.24272/j.issn.2095-8137.2019.032
Zhang, 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. Asian Herpetological Research, 8, 75–85.
https://doi.org/10.16373/j.cnki.ahr.160041