Abstract
In this report, we describe a new species of the Craugastor podiciferus species group from the premontane forest of the Pacific versant along the Costa Rican-Panamanian border. Mitochondrial DNA and karyotype analyses previously showed a marked genetic divergence between populations of the premontane forest of the Fila Costeña and the lowlands South Pacific Costa Rica near Panama. Analyses of the mitochondrial DNA sequences and the morphological variation revealed significant differences between the populations of the premontane forest relative to the other populations of C. stejnegerianus, including the type locality. We recognize these premontane populations as a new species and show that they differ from the typical C. stejnegerianus in the coloration of the venter, the head and the body proportions, and mtDNA divergence. With the addition of this new species, the C. podiciferus species group now contains nine species.
References
AmphibiaWeb (2016) AmphibiaWeb: Information on Amphibian Biology and Conservation [web application]. Berkeley, California. Available from: http://amphibiaweb.org (accessed 11 January 2016)
Barbour, T. (1926) New Amphibia. Occasional Papers of the Boston Society of Natural History, 5, 191–194.
Bickford, D., Lohman, D.J., Sodhi, N.S., Ng, P.K.L., Meier, R., Winker, K., Ingram, K.K. & Das, I. (2007) Cryptic species as a window on diversity and conservation. Trends in Ecology & Evolution, 22, 148–155.
http://dx.doi.org/10.1016/j.tree.2006.11.004Boulenger, G.A. (1896) Descriptions of new batrachians collected by Mr. C. G. Underwood in Costa Rica. Annals and Magazine of Natural History, Series 6, 18, 340–342.
http://dx.doi.org/10.1080/00222939608680464Chen, S.H. (2001) Cytogenetic study of the lower Central American frogs of the subgenus Craugastor (Anura: Leptodactylidae: Eleutherodactylus). Ph.D. dissertation, University of Miami, Florida, 369 pp.
Chen, S.H. (2005) Chromosomal variation in the rhodopis group of the southern Central American Eleutherodactyline frogs (Leptodactylidae: Eleutherodactylus). In: Donnelly, M.A., Crother, B.I., Guyer, C., Wake, M.H. & White, M.E. (Eds.), Ecology and evolution in the tropics: a herpetological perspective. The University of Chicago Press, Illinois, pp. 102–144.
Cope, E.D. (1886) Thirteenth contribution to the herpetology of tropical America. Proceedings of the American Philosophical Society, 23, 271–287.
Cope, E.D. (1893) Second addition to the knowledge of the Batrachia and Reptilia of Costa Rica. Proceedings of the American Philosophical Society, 31, 333–347.
Crawford, A.J. (2003) Huge populations and old species of Costa Rican and Panamanian dirt frogs inferred from mitochondrial and nuclear gene sequences. Molecular Ecology, 12, 2525–2540.
http://dx.doi.org/10.1046/j.1365-294X.2003.01910.xCrawford, A.J. & Smith, E.N. (2005) Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution, 35, 536–555.
http://dx.doi.org/10.1016/j.ympev.2005.03.006Crawford, A.J., Bermingham, E. & Polanía, C. (2007) The role of tropical dry forest as a long-term barrier to dispersal: a comparative phylogeographical analysis of dry forest tolerant and intolerant frogs. Molecular Ecology, 16, 4789–4807.
http://dx.doi.org/10.1111/j.1365-294X.2007.03524.xde Queiroz, K. (2007) Species concepts and species delimitation. Systematic Biology, 56, 879–886.
http://dx.doi.org/10.1080/10635150701701083Duellman, W.E. & Lehr, E. (2009) Terrestrial-breeding frogs (Strabomantidae) in Peru. Natur- und Tier-Verlag, Münster, 382 pp.
Edgar, R.C. (2004) MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research, 32, 1792–1797.
http://dx.doi.org/10.1093/nar/gkh340Fouquet, 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.
http://dx.doi.org/10.1371/journal.pone.0001109Frost, 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., 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.
http://dx.doi.org/10.1206/0003-0090(2006)297[0001:TATOL]2.0.CO;2Günther, A.C.L.G. (1885–1902) Biologia Centrali-Americana. Reptilia and Batrachia. Porter, London, 326 pp.
Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182.
Heinicke, M.P., Duellman, W.E. & Hedges, S.B. (2007) Major Caribbean and Central American frog faunas originated by ancient oceanic dispersal. Proceedings of the National Academy of Sciences, USA, 104, 10092–10097.
http://dx.doi.org/10.1073/pnas.0611051104Heinicke, M.P., Duellman, W.E., Trueb, L., Means, D.B., MacCulloch, R.D. & Hedges, S.B. (2009) A new frog family (Anura, Terrarana) from South America and an expanded direct–developing clade revealed by molecular phylogeny. Zootaxa, 2211, 1–35.
Hertz, A., Hauenschild, F., Lotzkat, S. & Köhler, G. (2012) A new golden frog species of the genus Diasporus (Amphibia, Eleutherodactylidae) from the Cordillera Central, western Panama. ZooKeys, 196, 23–46.
http://dx.doi.org/10.3897/zookeys.196.2774Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics, 17, 754–755.
http://dx.doi.org/10.1093/bioinformatics/17.8.754Lynch, J.D. (1980) A new frog of the genus Eleutherodactylus from western Panama. Transactions of the Kansas Academy of Science, 83, 101–105.
http://dx.doi.org/10.2307/3627802Lynch, J.D. (1986) The definition of the Middle American clade of Eleutherodactylus based on jaw musculature (Amphibia, Leptodactylidae). Herpetologica, 42, 248–258.
MacMillan, I., Gans, P.B. & Alvarado, G. (2004) Middle Miocene to present plate tectonic history of the southern Central American Volcanic Arc. Tectonophysics, 392, 325–348.
http://dx.doi.org/10.1016/j.tecto.2004.04.014McCranie, J.R. (2015) A checklist of the amphibians and reptiles of Honduras, with additions, comments on taxonomy, some recent taxonomic decisions, and areas of further studies needed. Zootaxa, 3931 (3), 352–386.
http://dx.doi.org/10.11646/zootaxa.3931.3.2Miyamoto, M.M. (1983) Biochemical variation in the frog Eleutherodactylus bransfordii: geographic patterns and cryptic species. Systematic Zoology, 32, 43‒51.
http://dx.doi.org/10.2307/2413219Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Center, Uppsala University, Uppsala, Sweden.
Padial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16.
http://dx.doi.org/10.1186/1742-9994-7-16Padial, J.M., Grant, T. & Frost, D.R. (2014) Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria. Zootaxa, 3825 (1), 1–132.
http://dx.doi.org/10.11646/zootaxa.3825.1.1Palumbi, S., Martin, A., Romano, S., McMillan, W.O., Stice, L. & Grabowski, G. (1991) The simple fool’s guide to PCR, version 2.0. Special Publication, Department of Zoology and Kewalo Marine Laboratory, University of Hawaii, Honolulu, Hawaii, USA.
Pérez-Ponce de León, G & Nadler, S.A. (2010) What we don't recognize can hurt us: A plea for awareness about cryptic species. Journal of Parasitology, 96, 453–464.
http://dx.doi.org/10.1645/GE-2260.1Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583.
http://dx.doi.org/10.1016/j.ympev.2011.06.012Rambaut, A., Suchard, M.A., Xie, W. & Drummond, A.J. (2014) Tracer v1.6.1. Available from: http://beast.bio.ed.ac.uk/Tracer (accessed 13 June 2016)
Santos-Barrera, G., Pacheco, J., Mendoza-Quijano, F., Bolaños, F., Chaves, G., Daily, G.C., Ehrlich, P.R. & Ceballos, G. (2008) Diversity, natural history and conservation of amphibians and reptiles from the San Vito Region, southwestern Costa Rica. Revista de Biología Tropical, 56, 755–778.
Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between two Continents, between two Seas. The University of Chicago Press, Illinois, 934 pp.
Savage, J.M. & Emerson, S.B. (1970). Central American frogs allied to Eleutherodactylus bransfordii (Cope): a problem of polymorphism. Copeia, 1970, 623–644.
http://dx.doi.org/10.2307/1442305Savage, J.M., McCranie, J.R. & Espinal, M. (1996) A new species of Eleutherodactylus from Honduras related to Eleutherodactylus bransfordii (Anura: Leptodactylidae). Proceedings of the Biological Society of Washington, 109, 366–372.
Scott, N.J. Jr. (1976) The abundance and diversity of the herpetofaunas of tropical forest litter. Biotropica, 8, 41–58.
http://dx.doi.org/10.2307/2387818Smith, M.A., Poyarkov, N.A. Jr. & Hebert, P.D.N. (2008) CO1 DNA barcoding amphibians: Take the chance, meet the challenge. Molecular Ecology Resources, 8, 235–246.
http://dx.doi.org/10.1111/j.1471-8286.2007.01964.xStamatakis, A. (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
http://dx.doi.org/10.1093/bioinformatics/btu033Streicher, J.W., Crawford, A.J. & Edwards, C.W. (2009) Multilocus molecular phylogenetic analysis of the montane Craugastor podiciferus species complex (Anura: Craugastoridae) in Isthmian Central America. Molecular Phylogenetics and Evolution, 53, 620–630.
http://dx.doi.org/10.1016/j.ympev.2009.07.011Tamura, K. (1992) Estimation of the number of nucleotide substitutions when there are strong transition–transversion and G + C–content biases. Molecular Biology and Evolution, 9, 678–687.
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.
http://dx.doi.org/10.1093/molbev/mst197Taylor, E.H. (1952) A review of the frogs and toads of Costa Rica. The University of Kansas Science Bulletin, 35, 577‒942.
http://dx.doi.org/10.5962/bhl.part.4328Vences, M., Thomas, M., Bonett, R.M. & Vieites, D.R. (2005) Deciphering amphibian diversity through DNA barcoding: Chances and challenges. Philosophical Transactions of the Royal Society B: Biological Sciences, 360, 1859–1868.
http://dx.doi.org/10.1098/rstb.2005.1717Wake, D.B. (2009) What salamanders have taught us about evolution. Annual Review of Ecology, Evolution, and Systematics, 40, 333–352.
http://dx.doi.org/10.1146/annurev.ecolsys.39.110707.173552