Open Access Open Access  Restricted Access Subscription Access

A new species of Andinobates (Amphibia: Anura: Dendrobatidae) from west central Panama



Dendrobatid frogs are among the best known anurans in the world, mainly due to their toxicity and associated bright colors. A recently described dendrobatid genus, Andinobates, comprises frogs distributed among the Colombian Andes and Panama. During field work in the Distrito de Donoso, Colón province, Panama, we found a poison frog that we here describe as a new species. The new species belongs to the A. minutus species group and is described herein as Andinobates geminisae sp. nov. This new species differs from all other members of the group by having uniformly orange smooth skin over the entire body and a distinctive male advertisement call. The new species is smaller than other colorful dendrobatids present in the area, such as Oophaga pumilio and O. vicentei. We also provide molecular phylogenetic analyses of mitochondrial DNA sequences of dendrobatids and summarize genetic distances among Andinobates species. Andinobates geminisae occurs in Caribbean versant rainforest on the westernmost edge of the known distribution of A. minutus, and represents the fourth species within this genus in Panama. This is vulnerable to habitat loss and excessive harvesting and requires immediate conservation plans to preserve this species with a restricted geographic range.



Andinobates geminisae sp. nov., conservation, DNA barcoding, geographic distribution, molecular phylogenetics, Panama, poison dart frogs

Full Text:

PDF/A (6.70 MB)


Akaike, H. (1974) A new look at the statistical model identification. IEEE Transactions on Automatic Control, 19, 716–723.

Altschul, S.F., Gish, W., Miller, W., Myers, E.W. & Lipman, D.J. (1990) Basic Local Alignment Search Tool. Journal of Molecular Biology, 215, 403–410.

Amézquita, A., Marquez, R., Medina, R., Mejia-Vargas, D., Kahn, T.R., Suarez, G. & Mazariegos, L. (2013) A new species of Andean poison frog, Andinobates (Anura: Dendrobatidae) from the northwestern Andes of Colombia. Zootaxa, 1620, 163–178.

Batista, A. & Köhler, G. (2008) Variation in Oophaga pumilio (Amphibia: Anura: Dendrobatidae) in western Panama. Salamandra, 44, 225234.

Bauer, L. (1986) A new genus and a new specific name in the dart poison frog family (Dendrobatidae, Anura, Amphibia). Ripa. Netherlands November, 1–12.

Bauer, L. (1988) Pijlgifkikkers en verwanten: de familie Dendrobatidae. Het Paludarium, 1–6.

Bauer, L. (1994) New names in the family Dendrobatidae (Anura, Amphibia). RIPA Fall, 1–6.

Boulenger, G.A. (1884 "1883") On a collection of frogs from Yurimaguas, Huallaga River, Northern Peru. Proceedings of the Zoological Society of London, 1883, 635–638.

Brown, J.L., Schulte, R. & Summers, K. (2006) A new species of Dendrobates (Anura : Dendrobatidae) from the Amazonian lowlands in Peru. Zootaxa, 1152, 45–58.

Brown, J.L., Twomey, E., Pepper, M. & Rodriguez, M.S. (2008) Revision of the Ranitomeya fantastica species complex with description of two new species from Central Peru (Anura : Dendrobatidae). Zootaxa, 1823, 1–24.

Brown, J.L., Twomey, E., Amézquita, A., Barbosa De Souza, M., Caldwell, J.P., Lötters, S., Von May, R., Melo-Sampaio, P.R., Mejía- Vargas, D., Perez-Peña, P., Pepper, M., Poelman, E.H., Sanchez-Rodriguez, M. & Summers, K. (2011) A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae). Zootaxa, 3083, 1–120.

Charif, R.A., Clark, C.W. & Fristrup, K.M. (2004) Raven 1.3 User’s Manual. Cornell Laboratory of Ornithology, Ithaca, NY, 167 pp.

Clough, M. & Summers, K. (2000) Phylogenetic systematics and biogeography of the poison frogs: evidence from mitochondrial DNA sequences. Biological Journal of the Linnean Society, 70, 515–540.

Collins, R., Boykin, L., Cruickshank, R. & Armstrong, K. (2011) Barcoding's next top model: an evaluation of nucleotide substitution models for specimen identification. Methods in Ecology and Evolution, 3, 457–465.

Crawford, A.J., Lips, K.R. & Bermingham, E. (2010) Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama. Proceedings of the National Academy of Sciences of the United States of America, 107, 13777–13782.

Daly, J.W., Myers, C.W. & Whittaker, N. (1987) Further classification on skin alkaloids from neotropical posion frogs (Dendrobatidae), with a general survey of toxic/noxious substances in the amphibia. Toxicon, 25, 1023–1095.

Duellman, W.E. & Trueb, L. (1994) Biology of Amphibians. The Johns Hopkins University Press, Baltimore and London, 670 pp.

Duméril, A.M.C. & Bibron, G. (1841) Erpétologie Genérale ou Histoire Naturelle Complète des Reptiles. Volume 8. Librarie Enclyclopedique de Roret, Paris, 784 pp.

ESRI (Environmental Systems Resource Institute) (2009) ArcMap 9.2. ESRI, Redlands, California.

Faivovich, J., Haddad, C.F.B., Garcia, P.C.A., Frost, D.R., Campbell, J.A. & Wheeler, W.C. (2005) Systematic review of the frog family Hylidae, with special reference to Hylinae: phylogenetic analysis and taxonomic revision. Bulletin of the American Museum of Natural History, 294, 1–240.[0001:SROTFF]2.0.CO;2

Felsenstein, J. (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Journal of Molecular Evolution, 17, 368–376.

Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783–791.

Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.

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. (2013) Amphibian Species of the World: an Online Reference. Version 5.6. American Museum of Natural History, New York, USA. Electronic Database. Available from: (accessed 9 January 2013)

Gorzula, S. (1996) The trade in dendrobatid frogs from 1987 to 1993. Herpetological Review, 27, 116–123.

Grant, T., Frost, D.R., Caldwell, J.P., Gagliardo, R., Haddad, C.F.B., Kok, P.J.R., Means, D.B., Noonan, B.P., Schargel, W.E. & Wheeler, W. (2006) Phylogenetic systematics of dart-poison frogs and their relatives (Amphibia, Athesphatanura, Dendrobatidae). Bulletin of the American Museum of Natural History, 299, 1–262.[1:PSODFA]2.0.CO;2

Hasegawa, M., Kishino, H. & Yano, T-a. (1987) Man’s place in Hominoidea as inferred from molecular clocks of DNA. Journal of Molecular Evolution, 26, 132–147.

Hebert, P.D.N., Ratnasingham, S. & de Waard, J.R. (2003) Barcoding animal life: cytochrome c oxidase subunit 1 divergences among closely related species. Proceedings of the Royal Society of London, Series B (Biological Sciences), 270, S96–S99.

Hof, C., Araujo, M.B., Jetz, W. & Rahbek, C. (2011) Additive threats from patho-gens, climate and land-use change for global amphibian diversity. Nature, 480, 516–519.

Jungfer, K.H., Lötters, L. & Jörgens, D. (2000) Der kleinste Pfeilgiftfrosch – eine neue Dendrobates-Art aus West-Panama. Herpetofauna, 22, 11–18.

Jungfer, K.H, Weygoldt, P. & Juraske, N. (1996) Dendrobates vicentei, ein neuer Pfeilgiftfrosch aus Zentral-Panama. Herpetofauna, 18, 17–26.

Kessing, B., Croom, H., Martin, A., McIntosh, C., Owen, M.W. & Palumbi, S.P. (2004) The Simple Fool’s Guide to PCR. Version 1.0. Special Publication. Dept. of Zoology, University of Hawaii, Honolulu, USA, 23 pp.

Köhler, G. (2012) Color Catalogue for Field Biologists. Herpeton, Offenbach, Germany, 49 pp.

Lassmann, T. & Sonnhammer, E.L. (2005) Kalign - an accurate and fast multiple sequence alignment algorithm. BMC Bioinformatics, 6, 298.

Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Carey, C., Livo, L., Pessier, A.P. & Collins, J.P. (2006) Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community. Proceedings of the National Academy of Sciences of the United States of America, 103, 3165–3170.

Littlejohn, M.J. (1977) Long-range acoustic communication in anurans: an integrated and evolutionary approach. In: Taylor, D.H. & Guttman, S.I. (Eds.), The reproductive biology of amphibians, Plenum Press, New York, pp. 263–294.

Lötters, S., Jungfer, K., Henkel, F.W. & Schmidt, W. (2007) Poison Frogs. Biology, Species & Captive Husbandry. Edition Chimaira, Frankfurt am Main, 668 pp.

Maddison, W.P. & Maddison, D.R. (2009) Mesquite: a modular system for evolutionary analysis. Version 2.6. Available from: (accessed 3 September 2014)

Meyer, C.P., Geller, J.B. & Paulay, G. (2005) Fine scale endemism on coral reefs: Archipelagic differentiation in turbinid gastropods. Evolution, 59, 113–125.

Myers, C.W. & Daly, J. (1976) Preliminary evaluation of skin toxins and vocalizations in taxonomic and evolutionary studies of poison-dart frogs (Dendrobatidae). Bulletin of the American Museum of Natural History, 157, 173–262.

Myers, C.W. & Daly, J. (1976) A new species of poison frog (Dendrobates) from Andean Ecuador, including an analysis of its skin toxins. Occasional Papers of the Museum of Natural History The University of Kansas Lawrence, Kansas, 59, 1–12.

Myers, C.W. (1982) Spotted poison frogs: Description of Three new Dendrobates from Western Amazonia, and resurrection of a lost species from "Chiriqui". American Museum Novitates, 2721, 23.

Narins, P.M., Lewis, E.R. & McClelland, B.E. (2000) Hyperextended call note repertoire of the endemic Madagascar treefrog Boophis madagascariensis (Rhacophoridae). Journal of Zoology (London), 250, 283–298.

Padial, J., Miralles, A., De la Riva, I. & Vences, M. (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16.

Pepper, M.P., Twomey, E. & Brown, J.L. (2007) The smuggling crisis. Leaf Litter Magazine, Tree Walkers International, 1.1, 5–7.

Posada, D. (2008) jModelTest: phylogenetic model averaging. Molecular Biology and Evolution, 25, 1253–1256.

Ratnasingham, S. & Hebert, P.D.N. (2007) BoLD: The barcode of life data system ( Molecular Ecology Notes, 7, 355–364.

Rivero, J.A. (1971) Un nuevo e interesante Dendrobates (Amphibia, Salientia) de Cerro Yapacana de Venezuela. Kasmera, 3, 389–396.

Roberts, J.L., Brown, J.L., von May, R., Arizabal, W., Presar, A., Symula, R., Schulte, R. & Summers, K. (2006) Phylogenetic relationships among poison frogs of the genus Dendrobates (Dendrobatidae): A molecular perspective from increased taxon sampling. Herpetological Journal, 16, 377–385.

Ryan, M.J., Savage, J.M., Lips, K.R. & Giermakowski, J.T. (2010) A new species of the Craugastor rugulosus series (Anura: Craugastoridae) from west-central Panama. Copeia, 2010, 405–409.

Saitou, N. & Nei, M. (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. Molecular Biology and Evolution, 4, 406–425.

Santos, J.C., Coloma, L.A. & Cannatella, D.C. (2003) Multiple, recurring origins of aposematism and diet specialization in poison frogs. Proceedings of the National Academy of Sciences of the United States of America, 100, 12792–12797.

Santos, J.C., Coloma, L.A., Summers, K., Caldwell, J.P., Ree, R. & Cannatella, D.C. (2009) Amazonian Amphibian Diversity Is Primarily Derived from Late Miocene Andean Lineages. PLoS Biol, 7, e1000056.

Santos, J.C. & Cannatella, D.C. (2011) Phenotypic integration emerges from aposematism and scale in poison frogs. Proceedings of the National Academy of Sciences of the United States of America, 108, 6175–6180.

Savage, J.M. (1968) The dendrobatid frogs of Central America. Copeia, 1968, 745–776.

Schmidt, O. (1858) Deliciae herpetologicae Musei Zoologici Cracoviensis, beschreibung der im K.K. Museum zu Krakau befindlichen, von J.V. Warszewitz in Neu-Granada und Bolivia Gesammelten. Ungeschwäntzen Batrachier. Denkschriften. Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klass, 14, 237–258.

Shreve, B. (1935) On a new Teiid and Amphibia from Panama, Ecuador, and Paraguay. Occasional Papers of the Boston Society of Natural History, 8, 209–218.

Silverstone, P.A. (1975) A revision of the poison–arrow frogs of the genus Dendrobates Wagler. Natural History Museum of Los Angeles County Science Bulletin, 21, 1–55.Swofford, D.L. (2004) PAUP*: Phylogenetic Analysis Using Parsimony (* and OtherMethods), Version 4.0b. Sunderland, MA: Sinauer Associates.

Symula, R., Schulte, R. & Summers, K. (2001) Molecular phylogenetic evidence for a mimetic radiation in Peruvian poison frogs supports a Mullerian mimicry hypothesis. Proceedings of the Royal Society of London Series B-Biological Science, 268, 2415–2421.

Sukumaran, J. & Holder, M.T. (2010) DendroPy: A Python library for phylogenetic computing. Bioinformatics, 26, 1569–1571.

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.

Tavaré, S. (1986) Some probabilistic and statistical problems on the analysis of DNA sequences. Lectures in Mathematics and the Life Sciences, 17, 57–86.

Twomey, E. & Brown J.L. (2008) Spotted Poison Frogs: rediscovery of a lost species and a new genus (Anura: Dendrobatidae) from northwestern Peru. Herpetologica, 64: 121–137.

Twomey, E., Brown, J.L., Amézquita, A. & Mejía-Vargas, D. (2011) Andinobates gen. nov. In: Brown, J.L., Twomey, E., Amézquita, A., Barbosa De Souza, M., Caldwell, J.P., Lötters, S., Von May, R., Melo-Sampaio, P.R., Mejía-Vargas, D., Perez-Peña, P., Pepper, M., Poelman, E.H., Sanchez-Rodriguez, M. & Summers, K. (2011) A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae). Zootaxa, 3083, 1–120.

Vences, M., Kosuch, J., Lötters, S., Widmer, A., Jungfer, K.H., Kohler, J. & Veith, M. (2000) Phylogeny and classification of poison frogs (Amphibia: dendrobatidae), based on mitochondrial 16S and 12S ribosomal RNA gene sequences. Molecular Phylogenetics and Evolution, 15, 34–40.

Wagler, J. (1830) Natürliches System der Amphibien, mit vorangehender Classification der Säugthiere und Vogel. Ein Beitrag zur vergleichenden Zoologie. J. G. Cotta, München, Stuttgart and Tübingen, 354 pp.

Wake, D.B. & Vredenburg, V.T. (2008) Are we in the midst of the sixth mass extinction? A view from the world of amphibians. Proceedings of the National Academy of Sciences of the United States of America, 105, 11466–11473.

Wang, I.J. (2011) Inversely related aposematic traits: reduced conspicuousness evolves with increased toxicity in a polymorphic poison-dart frog. Evolution, 65, 1637–1649.

Wang, I.J. & Shaffer, H.B. (2008) Rapid color evolution in an aposematic species: a phylogenetic analysis of color variation in the strikingly polymorphic strawberry poison-dart frog. Evolution, 62, 2742–2759.

Werle, E., Schneider, C., Renner, M., Volker, M. & Fiehn, W. (1994) Convenient single-step, one tube purification of PCR products for direct sequencing. Nucleic Acids Research, 22, 4354–4355.

Wollenberg, K.C., Veith, M., Noonan, B.P. & Lötters, S. (2006) Polymorphism versus species richness—systematics of large Dendrobates from the eastern Guiana Shield (Amphibia: Dendrobatidae). Copeia, 2006, 623–629.[623:PVSROL]2.0.CO;2

WWF, Hogan, M.C. & McGinley, M. (2010) "Isthmian-Atlantic moist forests". In: Encyclopedia of Earth. Eds. Cutler J. Cleveland (Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment). [First published in the Encyclopedia of Earth January 24, 2010. Last revised Date October 11, 2011. Available from: (accessed 4 March 2013)

Yang, Z. (1994) Maximum likelihood phylogenetic estimation from DNA sequences with variable rates over sites: approximate methods. Journal of Molecular Evolution, 39, 306–324.

Zwickl, D.J. (2006) Genetic Algorithm Approaches for the Phylogenetic Analysis of Large Biological Sequence Datasets under the Maximum Likelihood Criterion. Ph.D. Dissertation. The University of Texas, Austin, 125 pp.


  • There are currently no refbacks.

ISSN 1175-5326 (Print Edition) & ISSN 1175-5334 (Online Edition)
Published by Magnolia Press, Auckland, New Zealand
A founding journal of Biotaxa