Abstract
The Rufescent Burrowing Frog, Fejervarya rufescens, is thought to have a wide distribution across the Western Ghats in Peninsular India. This locally abundant but secretive species has a short breeding period, making it a challenging subject for field studies. We sampled 16 populations of frogs morphologically similar to F. rufescens in order to understand the variation among populations found across the Western Ghats. Our study shows significant morphological and genetic differences among the sampled populations, suggesting that F. ‘rufescens’ is a complex of several undescribed species. Using evidence from morphology and genetics, we confirm the presence of five distinct species in this group and formally describe four as new. The new species were delineated using a phylogeny based on three mitochondrial genes (16S, COI and Cytb) and a haplotype network of a nuclear gene (Rag1). Hereafter, the distribution of F. rufescens is restricted to the state of Karnataka and adjoining regions of northern Kerala. Three new species (Fejervarya kadar sp. nov., Fejervarya manoharani sp. nov. and Fejervarya neilcoxi sp. nov.) are from regions south of Palghat gap in the state of Kerala, and one (Fejervarya cepfi sp. nov.) from the northern Western Ghats state of Maharashtra. These findings indicate that Fejervarya frogs of the Western Ghats are more diverse than currently known. Our results will also have implications on the conservation status of F. rufescens, which was previously categorized as Least Concern based on its presumed wide geographical distribution. Furthermore, in order to facilitate a better taxonomic understanding of this region’s fejervaryan frogs, we divide all the known Fejarvarya species of the Western Ghats into four major groups—Fejervarya nilagirica group, Fejervarya rufescens group, Fejervarya sahyadris group and Fejervarya syhadrensis group, based on their morphological affinities.
References
AmphibiaWeb (2016) AmphibiaWeb: information on amphibian biology and conservation, University of California, Berkeley, California. Available from: http://www.amphibiaweb.org/ (accessed 16 November 2016)
Angulo, A. & Icochea, J. (2010) Cryptic species complexes, widespread species and conservation: lessons from Amazonian frogs of the Leptodactylus marmoratus group (Anura: Leptodactylidae). Systematics and Biodiversity, 8 (3), 357–370.
https://doi.org/10.1080/14772000.2010.507264
Bee, M.A., Suyesh, R. & Biju, S.D. (2013a) 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
Bee, M.A., Suyesh, R. & Biju, S.D. (2013b) The vocal repertoire of Pseudophilautus kani, a shrub frog (Anura: Rhacophoridae) from the Western Ghats of India. Bioacoustics, 22 (1), 67–85.
https://doi.org/10.1080/09524622.2012.712750
Biju, S.D. (2001) A synopsis to the frog fauna of the Western Ghats, India. ISBC - Occasional Publication, 1, 1–24.
Biju, 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/nature02019
Biju, S.D. & Bossuyt, F. (2005) Two new Philautus (Anura: Ranidae: Rhacophorinae) from Ponmudi Hill in the Western Ghats of India. Copeia, 2005 (1), 29–37.
https://doi.org/10.1643/CH-04-194R1
Biju, S.D., Dutta, S. & Padhye, A. (2004) Fejervarya rufescens. The IUCN Red List of Threatened Species 2004: e.T58288A91237431. Available from: https://doi.org/10.2305/IUCN.UK.2016-1.RLTS.T58288A91237431.en (accessed 15 September 2016)
Biju, S.D., Van Bocxlaer, I., Mahony, S., Dinesh, K.P., Radhakrishnan, C., Zachariah, A., Giri, V. & Bossuyt, F. (2011) A taxonomic review of the Night Frog genus Nyctibatrachus Boulenger, 1882 in the Western Ghats, India (Anura: Nyctibatrachidae) with description of twelve new species. Zootaxa, 3029, 1–96.
Biju, S.D., Garg, S., Gururaja, K.V., Shouche, Y. & Walujkar, S.A. (2014a) 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 (1), 37–123.
https://doi.org/10.4038/cjsbs.v43i1.6850
Biju, S.D., Garg, S., Mahony, S., Wijayathilaka, N., Senevirathne, G. & Meegaskumbura, M. (2014b) 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.
http://www.ctoz.nl/vol83/nr04/a04
Blair, C., Davy, C.M., Ngo, A., Orlov, N.L., Shi, H.T., Lu, S.Q., Gao, L., Rao, D.Q. & Murphy, R.W. (2013) Genealogy and demographic history of a widespread amphibian throughout Indochina. Journal of Heredity, 104 (1), 72–85.
https://doi.org/10.1093/jhered/ess079
Bossuyt, F., Meegaskumbura, M., Beenaerts, N., Gower, D.J., Pethiyagoda, R., Roelants, K., Mannaert, A., Wilkinson, M., Bahir, M.M., Manamendra-Arachchi, K., Ng, P.K., Schneider, C.J., Oommen, O.V. & Milinkovitch, M.C. (2004) Local endemism within the Western Ghats-Sri Lanka biodiversity hotspot. Science, 306 (5695), 479–481.
https://doi.org/10.1126/science.1100167
Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid frogs reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences, 97 (12), 6585–6590.
https://doi.org/10.1073/pnas.97.12.6585
Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Second edition. Taylor and Francis, London, 503 pp.
Boulenger, G.A. (1920) A monograph of the South Asian, Papuan, Melanesian and Australian frogs of the genus Rana. Records of the Indian Museum, 20, 1–226.
https://doi.org/10.5962/bhl.title.12471Chanda, S.K. (2002) Handbook—Indian Amphibians. Zoological Survey of India, Calcutta, 335 pp.
Charif, R.A., Waack, A.M. & Strickman, L.M. (2010) Raven Pro 1.4 User’s Manual. Cornell Lab of Ornithology, Ithaca, 367 pp.
Che, J., Chen, H.M., Yang, J.X., 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, 247–258.
https://doi.org/10.1111/j.1755-0998.2011.03090.x
Che, J., Zhou, W.W., Hu, J.S., Yan, F., Papenfuss, T.J., Wake, D.B. & Zhang, Y.P. (2010) Spiny frogs (Paini) illuminate the history of the Himalayan region and Southeast Asia. Proceedings of the National Academy of Sciences, 107 (31), 13765–13770.
https://doi.org/10.1073/pnas.1008415107
Daniels, R.J.R. (1992) Geographical Distribution Patterns of Amphibians in the Western Ghats, India. Journal of Biogeography, 19 (5), 521–529.
https://doi.org/10.2307/2845771
Daniels, R.J.R. (2005) Amphibians of Peninsular India. Indian Academy of Sciences & University Press, Hyderabad, 268 pp.
Das, I. & Dutta, S.K. (1998) Checklist of the amphibians of India, with English common names. Hamadryad, 23, 63–68.
Dinesh, K.P., Vijayakumar, S.P., Channakeshavamurthy, B.H., Torsekar, V.R., Kulkarni, N.U. & Shanker, K. (2015) Systematic status of Fejervarya (Amphibia, Anura, Dicroglossidae) from South and SE Asia with the description of a new species from the Western Ghats of Peninsular India. Zootaxa, 3999 (1), 79–94.
https://doi.org/10.11646/zootaxa.3999.1.5
Djong, T.H., Matsui, M., Kuramoto, M., Nishioka, M. & Sumida, M. (2011) A new species of the Fejervarya limnocharis complex from Japan (Anura, Dicroglossidae). Zoological Science, 28 (12), 922–929.
https://doi.org/10.2108/zsj.28.922
Dubois, A. (1975) Un nouveau complexe d’espèces jumelles distinguées par le chant: les grenouilles du Nèpal voisines de Rana limnocharis Boie (Amphibiens, Anoures). Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences, Série D: Sciences Naturelles, 281, 1717–1720.
Dubois, A. (1984) Note preliminaire sur le groupe de Rana limnocharis Gravenhorst, 1829 (Amphibiens, Anoures). Alytes, 3 (4), 143–159.
Dubois, A. (1987 “1986”) Miscellanea taxinomica batrachologica (I). Alytes, 5, 7–95.
Dubois, A. (1999) South Asian Amphibia: a new frontier for taxonomists. Journal of South Asian Natural History, 4 (1), 1–11.
Dubois, A. & Ohler, A. (2000) Systematics of Fejervarya limnocharis (Gravenhorst, 1829) (Amphibia, Anura, Ranidae) and related species. 1. Nomenclatural status and type-specimens of the nominal species Rana limnocharis Gravenhorst, 1829. Alytes, 18 (1–2), 15–50.
Dubois, A., Ohler, A. & Biju, S.D. (2001) A new genus and species of Ranidae (Amphibia, Anura) from south-western India. Alytes, 19 (2–4), 53–79.
Dutta, S.K. (1997) Amphibians of India and Sri Lanka (Checklist and Bibliography). Odyssey Publishing House, Bubaneswar, xiii + 342 + xxii pp.
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 (6), 794–819.
https://doi.org/10.1093/sysbio/52.6.794
Fei, L., Ye, C.Y., Jiang, J.P. & Xie, F. (2002) On taxonomic status of Rana limnocharis group with revision of nomenclature of the rice frog from China. Herpetologica Sinica, 9, 88–96.
Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N.J. (2007a) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS ONE, 2 (10), e1109.
https://doi.org/10.1371/journal.pone.0001109
Fouquet, A., Vences, M., Salducci, M.D., Meyer, A., Marty, C., Blanc, M. & Gilles, A. (2007b) Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera species groups. Molecular Phylogenetics and Evolution, 43 (2), 567–582.
https://doi.org/10.1016/j.ympev.2006.12.006
Frost, D.R. (2016) Amphibian species of the world: an online reference. Version 6.0. American Museum of Natural History, New York, USA. Available from: http://research.amnh.org/herpetology/amphibia/index.html (accessed 16 November 2016)
Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, 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;2Funk, W.C., Caminer, M. & Ron, S.R. (2011) High levels of cryptic species diversity uncovered in Amazonian frogs. Proceedings of the Royal Society B, 279, 1806–1814.
https://doi.org/10.1098/rspb.2011.1653
Garg, S. & Biju, S.D. (2016) Molecular and morphological study of Leaping Frogs (Anura, Ranixalidae) with description of two new species. PloS ONE, 11, e0166326.
https://doi.org/10.1371/journal.pone.0166326
Garg, S., Suyesh, R., Sukeshan, S. & Biju, S.D. (2017) Seven new species of Night Frogs (Anura, Nyctibatrachidae) from the Western Ghats Biodiversity Hotspot of India, with remarkably high diversity of diminutive forms. PeerJ, 5, e3007.
https://doi.org/10.7717/peerj.3007
Grosjean, S. & Dubois, A. (2011) Description of the advertisement calls of nine species of Fejervarya Bolkay, 1915 and Minervarya Dubois, Ohler and Biju, 2001 from China, India and Nepal. Alytes, 27, 117–141.
Hasan, M., Islam, M.M., Khan, M.M.R., Alam, M.S., Kurabayashi, A., Igawa, T., Kuramoto, M. & Sumida, M. (2012) Cryptic anuran biodiversity in Bangladesh revealed by mitochondrial 16S rRNA gene sequences. Zoological science, 29 (3), 162–172.
https://doi.org/10.2108/zsj.29.162
Howlader, M.S.A. (2011) Cricket frog (Amphibia: Anura: Dicroglossidae): two regions of Asia are corresponding two groups. Bangladesh Wildlife Bulletin, 5, 1–7.
Howlader, M.S.A., Nair, A. & Merilä, J. (2016) A new species of frog (Anura: Dicroglossidae) discovered from the mega city of Dhaka. PLoS ONE, 11, e0149597.
https://doi.org/10.1371/journal.pone.0149597
Iskandar, D.T. (1998) The Amphibians of Java and Bali. Research and Development Centre for Biology. LIPI and GEF Biodiviersity Collections Project, Bogor, xix + 117 pp. + 26 pl.
Islam, M.M., Kurose, N., Khan, M.M.R., Nishizawa, T., Kuramoto, M., Alam, M.S., Hasan, M.K., Kurniawan, N., Nishioka, M. & Sumida, M. (2008) Genetic divergence and reproductive isolation in the genus Fejervarya (Amphibia: Anura) from Bangladesh inferred from morphological observations, crossing experiments, and molecular analyses. Zoological Science, 25 (11), 1084–1105.
https://doi.org/10.2108/zsj.25.1084
Joshy, S.H. & Kuramoto, M. (2008) Comparative chromosome studies of five species of the genus Fejervarya (Anura: Ranidae) from South India. Cytologia, 73 (3), 243–250.
https://doi.org/10.1508/cytologia.73.243
Jerdon, T.C. (1853) Catalogue of reptiles inhabiting the Peninsula of India. Journal of the Asiatic Society of Bengal, 22, 522–534.
Jerdon, T.C. (1870) Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, 1870, 66–85.
Kadadevaru, G.G., Kanamadi, R.D. & Schneider, H. (2000) Advertisement call of two Indian ranids, Indirana beddomii and Tomopterna rufescens. Amphibia-Reptilia, 21, 242–246.
Kadadevaru, G.G., Kanamadi, R.D. & Schneider, H. (2002) Advertisement call, courtship and mating behaviour of the frog, Limnonectes syhadrensis from Western Ghats. Current Science, 82, 503–505.
Kosuch, J., Vences, M., Dubois, A., Ohler, A. & Bo ̈ hme, W. (2001) Out of Asia: Mitochondrial DNA Evidence for an Oriental Origin of Tiger Frogs, Genus Hoplobatrachus. Molecular Phylogenetics and Evolution, 21 (3), 398–407.
https://doi.org/10.1006/mpev.2001.1034
Kotaki, M., Kurabayashi, A., Matsui, M., Khonsue, W., Djong, T.H., Tandon, M. & Sumida, M. (2008) Genetic divergences and phylogenetic relationships among the Fejervarya limnocharis complex in Thailand and neighboring countries revealed by mitochondrial and nuclear genes. Zoological Science, 25 (4), 381–390.
https://doi.org/10.2108/zsj.25.381
Kotaki, M., Kurabayashi, A., Matsui, M., Kuramoto, M., Djong, T.H. & Sumida, M. (2010) Molecular Phylogeny of the diversified frogs of genus Fejervarya (Anura: Dicroglossidae). Zoological Science, 27 (5), 386−395.
https://doi.org/10.2108/zsj.27.386
Kumar, K.S., Anoop, V.S., Sivakumar, K.C., Raghunathan, D., Manoj, P., Deuti, K. & George, S. (2017) An additional record of Fejervarya manoharani from the Western Ghats with a description of its complete mitochondrial genome. Zootaxa, 4277 (4), 491–502.
https://doi.org/10.11646/zootaxa.4277.4.2Kurabayashi, A., Kuramoto, M., Joshy, S.H. & Sumida, M. (2005) Molecular phylogeny of the ranid frogs from southwest India based on the mitochondrial ribosomal RNA gene sequences. Zoological Science, 22 (5), 525–534.
https://doi.org/10.2108/zsj.22.525
Kuramoto, M. & Dubois, A. (2009) Bioacoustic studies on three frog species from the Western Ghats, South India. Current Herpetology, 28, 65–70.
https://doi.org/10.3105/018.028.0203
Kuramoto, M. & Joshy, S.H. (2001) Advertisement call structures of frogs from Southwestern India, with some ecological and taxonomic notes. Current Herpetology, 20, 85–95.
https://doi.org/10.5358/hsj.20.85
Kuramoto, M., Joshy, S.H., Kurabayashi, A. & Sumida, M. (2008 “2007”) The genus Fejervarya (Anura: Ranidae) in central Western Ghats, India, with descriptions of four new cryptic species. Current Herpetology, 26, 81–105.
Librado, P. & Rozas, J. (2009) DnaSP v5: A software for comprehensive analysis of DNA polymorphism data. Bioinformatics, 25 (11), 1451–1452.
https://doi.org/10.1093/bioinformatics/btp187
Matsui, M., Toda, M. & Ota, H. (2008 “2007”) A new species of frog allied to Fejervarya limnocharis from southern Ryukyuas, Japan (Amphibia: Ranidae). Current Herpetology, 26, 65–79.
McLeod, D.S. (2010) Of least concern? Systematics of a cryptic species complex: Limnonectes kuhlii (Amphibia: Anura: Dicroglossidae). Molecular Phylogenetics and Evolution, 56 (3), 991–1000.
https://doi.org/10.1016/j.ympev.2010.04.004
Meenakshi, K., Suraj, T., Bhagwati, S.S., Sujith, V.G., Santhoshkumar, K. & Sanil, G. (2010) Molecular resolution of four Fejervarya Species from Western Ghats (India) with their intrageneric phylogeny based on COI, Cyt B, 12S and 16S rRNA Genes. Asian Journal of Experimental Biological Science, 1 (4), 782–786.
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.
Myers, N., Mittermeier, R.A., Mittermeier, C.G., Da Fonseca, G.A. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.
https://doi.org/10.1038/35002501
Ohler, A., Deuti, K., Grosjean, S., Paul, S., Ayyaswamy, A.K., Ahmed, M.F. & Dutta, S.K. (2009) Small-sized dicroglossids from India, with the description of a new species from West Bengal, India. Zootaxa, 2209, 43–56.
Ohler, A., Dutta, S.K. & Dubois, A. (2014) Morphological evolution in frogs of the genera Fejervarya, Minervarya, Sphaerotheca and Zakerana (Dicroglossidae). Pranikee-Journal of Zoological Society of Orissa, 16, 1–12.
Posada, D. & Crandall, K.A. (1998) Modeltest: testing the model of DNA substitution. Bioinformatics, 14 (9), 817–818.
https://doi.org/10.1093/bioinformatics/14.9.817
Purkayastha, J. & Matsui, M. (2012) A new species of Fejervarya (Anura: Dicroglossidae) from Mawphlang, northeastern India. Asian Herpetological Research, 3 (1), 31–37.
https://doi.org/10.3724/sp.j.1245.2012.00031
Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of advanced frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61 (2), 543–583.
https://doi.org/10.1016/j.ympev.2011.06.012
Rao, C.R.N. (1920) Some South Indian batrachians. Journal of the Bombay Natural History Society, 27, 119–127.
Rao, C.R.N. (1922) Notes on Batrachia. Journal of the Bombay Natural History Society, 28, 439–447.
Rao, C.R.N. (1937) On some new forms of Batrachia from S. India. Proceedings of the Indian Academy of Sciences: Section B, 6, 387–427.
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/btg180Roy, D. & Elepfandt, A. (1993) Bioacoustic analysis of frog calls from northeast India. Journal of Biosciences, 18 (3), 381–393.
https://doi.org/10.1007/bf02702996Savage, 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.
Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12 (4), 335–337.
https://doi.org/10.1007/s13127-011-0056-0
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 Entomological Society of America, 87 (6), 651–701.
https://doi.org/10.1093/aesa/87.6.651
Stamatakis, A. (2006) RAxML-VI-HPC: Maximum Likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22 (21), 2688–2990.
https://doi.org/10.1093/bioinformatics/btl446
Stamatakis, A., Hoover, P., Rougemont, J. & Renner, S. (2008) A rapid bootstrap algorithm for the RAxML Web Servers. Systematic biology, 57 (5), 758–771.
https://doi.org/10.1080/10635150802429642
Stephens, M., Smith, N.J. & Donnelly, P. (2001) A new statistical method for haplotype reconstruction from population data. American Journal of Human Genetics, 68 (4), 978–989.
https://doi.org/10.1086/319501
Sumida, M., Kotaki, M., Islam, M.M., Djong, T.H., Igawa, T., Kondo, Y., Matsui, M., De Silva, A., Khonsue, W. & Nishioka, M. (2007) Evolutionary relationships and reproductive isolating mechanisms in the rice frog (Fejervarya limnocharis) species complex from Sri Lanka, Thailand, Taiwan and Japan, inferred from mtDNA gene sequences, allozymes, and crossing experiments. Zoological science, 24 (6), 547–562.
https://doi.org/10.2108/zsj.24.547
Swofford, D.L. (2002) PAUP*: Phylogenetic Analysis Using Parsimony (* and other methods). Version 4.0b10. Sinauer Association Inc., Sunderland, Massachusetts.
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
Van Bocxlaer, I., Roelants, K., Biju, S.D., Nagaraju, J. & Bossuyt, F. (2006) Late Cretaceous vicariance in Gondwanan amphibians. PloS ONE, 1 (1), e74.
https://doi.org/10.1371/journal.pone.0000074
Van Bocxlaer, I., Biju, S.D., Willaert, B., Giri, V.B., Shouche, Y.S. & Bossuyt, F. (2012) Mountain-associated clade endemism in an ancient frog family (Nyctibatrachidae) on the Indian subcontinent. Molecular Phylogenetics and Evolution, 62 (3), 839–847.
https://doi.org/10.1016/j.ympev.2011.11.027
Veith, M., Kosuch, J., Ohler, A. & Dubois, A. (2001) Systematics of Fejervarya limnocharis (Gravenhorst, 1829) (Amphibia, Anura, Ranidae) and related species. 2. Morphological and molecular variation in frogs from the Greater Sunda Islands (Sumatra, Java, Borneo) with the definition of two species. Alytes, 19 (1), 5–28.
Wiens, J.J., Sukumaran, J., Pyron, R.A. & Brown, R.M. (2009) Evolutionary and biogeographic origins of high tropical diversity in Old World frogs (Ranidae). Evolution, 63 (5), 1217–1231.
https://doi.org/10.1111/j.1558-5646.2009.00610.x