Abstract
Systematic relationships of fanged frogs usually associated with Limnonectes kuhlii are assessed using 15 samples from Japan, Chinese Mainland and Taiwan, Vietnam, Laos, Thailand, Malaysia (Borneo), and Indonesia. Phylogenetic relationship inferred from the mitochondrial 12S rRNA, tRNA val , and 16S rRNA gene sequences revealed that fanged frogs sampled are not monophyletic with the topotypic L. kuhlii from Java. Samples from Yunnan of southern China (L. bannaensis), northern Laos and central Vietnam, and those from Jiangxi of eastern China (L. fujianensis), Taiwan and Japan (L. namiyei), respectively, form monophyletic groups, and are collectively sister to the Thai sample (L. megastomias). All these samples, L. fragilis from Hainan of southern China, and a group of Bornean samples show unresolved relationships with Javanese L. kuhlii. From the resultant phylogeny and genetic distances found among samples, L. "kuhlii" from Taiwan and L. fujianensis, and L. "kuhlii" from northern Laos and central Vietnam and L. bannaensis, respectively, are surmised to be conspecific. These fanged frogs are morphologically similar to, but phylogenetically distant from, L. kuhlii sensu stricto. Limnonectes namiyei, L. fujianensis, and L. bannaensis are considered to have a common ancestor whose chromosome number was 2n=22, unlike L. fragilis, L. kuhlii and many other frogs with 2n=26 chromosomes.References
Alam, M.S., Igawa, T., Khan, Md.M.R., Islam, M.M., Kuramoto, M., Matsui, M., Kurabayashi, A. & Sumida, M. (2008) Genetic divergence and evolutionary relationships in six species of genera Hoplobatrachus and Euphlyctis (Amphibia: Anura) from Bangladesh and other Asian countries revealed by mitochondrial gene sequences. Molecular Phylogenetics and Evolution, 48, 515–527.
Bain, R. & Nguyen, Q.T. (2004) Herpetofaunal diversity of Ha Giang Province in northeastern Vietnam, with descriptions of two new species. American Museum Novitates, 3453, 1–42.
Boulenger, G.A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Musuem, 20, 1–223.
Che, J., Pang, J., Zhao, H., Wu, G.-F., Zhao, E.-M. & Zhang, Y.-P. (2007) Molecular phylogeny of the Chinese ranids inferred from nuclear and mitochondrial DNA sequences. Biochemical Systematics and Ecology, 35, 29–39.
Emerson, S.B., Inger, R.F. & Iskandar, D. (2000) Molecular systematics and biogeography of the fanged frogs of Southeast Asia. Molecular Phylogenetics and Evolution, 16, 131–142.
Evans, B.J., Brown, R.M., McGuire, J.A., Supriatna, J., Andayani, N., Diesmos, A., Iskandar, D., Melnick, D.J. & Cannatella, D.C. (2003) Phylogenetics of fanged frogs: Testing biogeographical hypotheses at the interface of the Asian and Australian faunal zones. Systematic Biology, 52, 794–819.
Farris, J.S., Kallersjo, M., Kluge, A.G. & Bult, C. (1994) Testing significance of incongruence. Cladistics, 10, 315–319.
Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.
Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N. J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS ONE, 2, e1109.
Frost, D.R. (2009) Amphibian Species of the World: an Online Reference, version 5.3 (12 February, 2009). Electronic database accessible at: http://research.amnh.org/ herpetology/amphibia/index.php, American Museum of Natural History, New York.
Hedges, S.B. (1994) Molecular evidence for the origin of birds. Proceedings of the National Academy of Sciences USA, 91, 2621–2624.
Hillis, D.M. & Bull, J.J. (1993) An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic Biology, 42, 182–192.
Hillis, D.M., Mable, B.K., Larson, A., Davis, S.K. & Zimmer, E.A. (1996) Nucleic acids IV: sequencing and cloning. In D.M. Hillis, C. Moritz & B.K. Mable (eds) Molecular Systematics, Sinauer Associates, Sunderland, Massachusetts: 321–378.
Huelsenbeck, J.P. & Rannala, B. (2004) Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models. Systematic Biology, 53, 904–913.
Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
Huelsenbeck, J.P., Ronquist, F., Nielsen, R. & Bollback, J.P. (2001) Bayesian inference of phylogeny and its impact on evolutionary biology. Science, 294, 2310–2314.
Inger, R.F. (1966) The systematics and zoogeography of the Amphibia of Borneo. Fieldiana: Zoology, 52, 1–402.
Inger, R.F. & Tan, F.-L. (1996) Checklist of the frogs of Borneo. Raffles Bulletin of Zoology, 44, 551–574.
Inger, R.F., Stuart, B.L. & Iskandar, D.T. (2009) Systematics of a widespread Southeast Asian frog, Rana chalconota (Amphibia: Anura: Ranidae). Zoological Journal of the Linnean Society, 155, 123–147.
Iskandar, D.T. (1998) The Amphibians of Java and Bali. Research and Development Centre for Biology-Lipi. 117 pp.
Joshy, S.H., Alam, M.S., Kurabayashi, A., Sumida, M. & Kuramoto, M. (2009) Two new species of the genus Euphlyctis (Anura: Ranidae) from southwestern India as revealed by molecular and morphological comparisons. Alytes, 26, 97–116.
Kou, Z.T., Li, W.H., Jin, A.L. & Song, Z. (1990) Rare karyotype of genus Rana- A preliminary study on the karyotype and C-banding pattern of Rana kuhlii Dumeril et Bibron. In E.-M. Zhao (ed) From Water onto Land, Chinese Forestry Publishing House, Beijing: 159–163.
Kuramoto, M. (1972) Karyotypes of the six species of frogs (genus Rana) endemic to the Ryukyu Islands. Caryologia, 25, 547–559.
Kuramoto, M. (1980) Karyotypes of several frogs from Korea, Taiwan and the Philippines. Experientia, 36, 826–828.
Leaché, A.D. & Reeder, T.W. (2002) Molecular systematics of the eastern fence lizard (Sceloporus undulatus): a comparison of parsimony, likelihood, and Bayesian approaches. Systematic Biology, 51, 44–68.
Li, S.S. (1991) The study of karyotype, C-banding and Ag-NORs of Rana kuhlii (Ranidae, Anura). Hereditas, 13, 2–23.
Lue, K.-Y., Tu, M.-C. & Hsiang, G. (1999) Atlas of Taiwan Amphibians and Reptililes. Nature Press, Taipei, 343 pp.
Maeda, N. & Matsui, M. (1989) Frogs and toads of Japan. Bun-ichi Sogo Shuppan Co., Ltd., Tokyo, 206 pp.
Matsui, M., Seto, T. & Utsunomiya, T. (1986) Acoustic and karyotypic evidence for specific separation of Polypedates megacephalus from P. leucomystax. Journal of Herpetology, 20, 483–489.
Matsui, M., Chan-Ard, T. & Nabhitabhata, J. (1996) Distinct specific status of Kalophrynus pleurostigma interlineatus (Anura, Microhylidae). Copeia, 1996, 440–445.
Matsui, M., Ito, H., Shimada, T., Ota, H., Saidapur, S.K., Khonsue, W., Tanaka-Ueno, T. & Wu, G. (2005) Taxonomic relationships within the pan-oriental narrow-mouth toad Microhyla ornata as revealed by mtDNA analysis (Amphibia, Anura, Microhylidae). Zoological Science, 22, 489–495.
Matsui, M., Maryati, M., Shimada, T. & Sudin, A. (2007a) Resurrection of Staurois parvus from S. tuberilinguis from Borneo (Amphibia, Ranidae). Zoological Science, 24, 101–106.
Matsui, M., Toda, M. & Ota, H. (2007b) A new species of frog allied to Fejervarya limnocharis from the Southern Ryukyus, Japan (Amphibia: Ranidae). Current Herpetology, 26, 65–79.
Matsui, M., Yambun, P. & Sudin, A. (2007c) Taxonomic relationships of Ansonia anotis Inger, Tan, and Yambun, 2001 and Pedostibes maculatus (Mocquard, 1890), with a description of a new genus (Amphibia, Bufonidae). Zoological Science, 24, 1159–1166.
Matsui, M., Tominaga, A., Liu, W.-Z., Khonsue, W., Grismer , L.L., Diesmos, A.C., Das, I., Sudin, A., Yambun, P., Yong, H.-S., Sukumaran, J. & Brown, R.M. (2009) Phylogenetic relationships of Ansonia from Southeast Asia inferred from mitochondrial DNA sequences: Systematic and biogeographic implications (Anura: Bufonidae). Molecular Phylogenetics and Evolution, (2010), doi:10.1016/j.ympev.2009.08.003
McLeod, D.S. (2008) A new species of big-headed, fanged dicroglossine frog (Genus Limnonectes) from Thailand. Zootaxa, 1807, 26–46.
Morescalchi, A. (1973) Amphibia. In A.B, Chiarelli & E. Capanna (eds) Cytotaxonomy and Vertebrate Evolution, Academic Press, New York & London: 233–348.
Nguyen, V.S., Ho, T.C. & Nguyen, Q.T. (2009) Herpetofauna of Vietnam. Edition Chimaira, Frankfurt am Main, 768 pp.
Nylander, J.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre, Uppsala Univ, Uppsala.
Peng, Q.L., Wang ,Y., Xu, J.C., Tang, X., Zhang, J.F., Ji, D.L., Hu, L.L. & Nie, L.W. (2005) Karyotype,C-band and Ag-NORs of Limnonectes fujianensis. Chinese Journal of Zoology, 40, 77–78.
Posada, D. & Crandall, K.A. (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics, 14, 817–818.
Rambaut, A. & Drummond, A.J. (2007) Tracer version 1.4. Available from: http://tree.bio.ed.ac.uk/software/tracer/.
Rannala, B. & Yang, Z. (1996) Probability distribution of molecular evolutionary trees: a new method of phylogenetic inference. Journal of Molecular Evolution, 43, 304–311.
Rodriguez, F., Oliver, J.F., Marin, A. & Medina, J.R. (1990) The general stochastic model of nucleotide substitution. Journal of Theoretical Biology, 142, 485–501.
Stejneger, L. (1907) Herpetology of Japan and adjacent territory. Bulletin of United States National Museum, 58, 1–577.
Swofford, D. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods), Version 4. Sinauer Associates, Sunderland, MA.
Tamura, K. & Nei, M. (1993) Estimation of the number of nucleotide substitutions in the control region of mitochondrial DNA in humans and chimpanzees. Molecular Biology and Evolution, 10, 512–526.
Thompson, J.D., Higgins, D.G. & Gibson, T.J. (2007) 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, 4673–4680.
Toda, M., Matsui, M., Nishida, M. & Ota, H. (1998) Genetic divergence among Southeast and East Asian populations of Rana limnocharis (Amphibia: Anura), with special reference to sympatric cryptic species in Java. Zoological Science, 15, 607–613.
Tschudi, J.J.V. (1838) Classification der Batrachier, mit Berucksichtigung der fossilen Thiere dieser Abtheilung der Reptilien. Mémoires de la Société des Sciences Naturelles de Neuchatel, 2, 1–99.
Tsuji, H. & Matsui, M. (2002) Male-male combat and head morphology in a fanged frog (Rana kuhlii) from Taiwan. Journal of Herpetology, 36, 520–526.
Wynn, A. & Heyer, W.R. (2001) Do geographically widespread species of tropical amphibians exist? An estimate of genetic relatedness within the neotropical frog Leptodactylus fuscus (Anura, Leptodactylidae). Tropical Zoology, 14, 255–285.
Xu, J.C., Tang, X.S., Peng, Q.L. & Nie, L.W. (2007) Study on karyotypes diversity of five different geographical population in Limnonects fujianensis. Journal of Biology, 24, 43–45.
Ye, C.Y. & Fei, L. (1994) A new species of family Ranidae- Limnonectes fujianensis from Fujian, China. Acta Zootaxonomica Sinica, 19, 494–499.
Ye, C.-Y., Fei, L., Xie, F. & Jiang, J.-P. (2007) A new Ranidae species from China-Limnonectes bannaensis (Ranidae: Anura). Zoological Research, 28, 545–550.
Zhang, J.-F., Nie, L.-W., Peng, Q.-L., Ge, Y.-D., Wang, Y., Xu, J.-C. & Tang, X.-S. (2005) Relationships among the Chinese group of Limnonectes based on mitochondrial 12S and 16S rRNA sequences. Acta Zoologica Sinica, 51, 354–359.