Open Access Open Access  Restricted Access Subscription or Fee Access


DOI: http://dx.doi.org/10.11646/zootaxa.4004.1.1

Phylogenetic systematics of egg-brooding frogs (Anura: Hemiphractidae) and the evolution of direct development

SANTIAGO CASTROVIEJO-FISHER, JOSÉ M. JR. PADIAL, IGNACIO DE LA RIVA, JOSÉ P. POMBAL, JR., HELIO R. DA SILVA, FERNANDO J. M. ROJAS-RUNJAIC, ESTEBAN MEDINA-MÉNDEZ, DARREL R. FROST

Abstract


Egg-brooding frogs (Hemiphractidae) are a group of 105 currently recognized Neotropical species, with a remarkable diversity of developmental modes, from direct development to free-living and exotrophic tadpoles. Females carry their eggs on the back and embryos have unique bell-shaped gills. We inferred the evolutionary relationships of these frogs and used the resulting phylogeny to review their taxonomy and test hypotheses on the evolution of developmental modes and bell-shaped gills. Our inferences relied on a total evidence parsimony analysis of DNA sequences of up to 20 mitochondrial and nuclear genes (analyzed under tree-alignment), and 51 phenotypic characters sampled for 83% of currently valid hemiphractid species. Our analyses rendered a well-resolved phylogeny, with both Hemiphractidae (sister of Athesphatanura) and its six recognized genera being monophyletic. We also inferred novel intergeneric relationships [((Cryptobatrachus, Flectonotus), (Stefania, (Fritziana, (Hemiphractus, Gastrotheca))))], the non-monophyly of all species groups previously proposed within Gastrotheca and Stefania, and the existence of several putative new species within Fritziana and Hemiphractus. Contrary to previous hypotheses, our results support the most recent common ancestor of hemiphractids as a direct-developer. Free-living aquatic tadpoles apparently evolved from direct-developing ancestors three to eight times. Embryos of the sister taxa Cryptobatrachus and Flectonotus share a pair of single gills derived from branchial arch I, while embryos of the clade including the other four genera have two pairs of gills derived from branchial arches I and II respectively. Furthermore, in Gastrotheca the fusion of the two pairs of gills is a putative synapomorphy. We propose a revised taxonomy concordant with our optimal topologies.

 


Keywords


Amphibia, Cryptobatrachus, Flectonotus, Fritziana, Gastrotheca, gills, Hemiphractus, Neotropics, parsimony, Stefania, taxonomy, total evidence, tree-alignment

Full Text:

PDF/A (4.18 MB)

References


Acuña J., F. & Lugli, L. (2009) Reproductive biology, vocalizations, and tadpole morphology of Rupirana cardosoi, an anuran with uncertain affinities. South American Journal of Herpetology, 4, 173–178.

http://dx.doi.org/10.2994/057.004.0208

Adamson, L., Harrison, R.G. & Bayley, I. (1960) The development of the whistling frog Eleutherodactylus martinicensis of Barbados. Proceedings of the Zoological Society of London, 133, 453–469.

http://dx.doi.org/10.1111/j.1469-7998.1960.tb05573.x

Aguayo, R. (2009) Nymphargus bejaranoi (Cannatella, 1980) Anura – Centrolenidae. In: Ministerio de Medio Ambiente y Agua (Ed.), Libro rojo de la fauna silvestre de vertebrados de Bolivia. La Paz, Bolivia, pp 171–172.

Aguilar, C., Siu-Ting, K. & Venegas, P. (2007) The rheophilous tadpole of Telmatobius atahualpai Wiens, 1993 (Anura: Ceratophryidae). South American Journal of Herpetology, 2, 165–174.

http://dx.doi.org/10.2994/1808-9798(2007)2[165:TRTOTA]2.0.CO;2

Altig, R. & Johnston, G.F. (1989) Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. Herpetological Monograph, 3, 81–109.

http://dx.doi.org/10.2307/1466987

Anganoy-Criollo, M. (2013) Tadpoles of the High-Andean Hyloxalus subpunctatus (Anura: Dendrobatidae) with description of larval variation and species distinction by larval morphology. Papéis Avulsos de Zoologia, 53, 211–224.

Austin, J.D., Lougheed, S.C., Tanner, K., Chek, A.A., Bogart, J.P. & Boag P.T. (2002) A molecular perspective on the evolutionary affinities of an enigmatic neotropical frog, Allophryne ruthveni. Zoological Journal of the Linnean Society, 134, 335–346.

http://dx.doi.org/10.1046/j.1096-3642.2002.00011.x

Banks C.B., Birkett, J.R., Dunn, R.W. & Martin, A.A. (1983) Development of Litoria infrafrenata (Anura: Hylidae). Transactions of the Royal Society of South Australia, 107, 197–200.

http://biodiversitylibrary.org/page/41128317

Barrio-Amorós, C.L. (1999 “1998”) Sistematica y biogeografía de los anfibios (Amphibia) de Venezuela. Acta Biologica Venezuelica, 18, 1–93.

Barrio-Amorós, C.L. & Fuentes-Ramos, O. (2003) A new species of Stefania (Anura: Hylidae: Hemiphractinae) from the summit of Cerro Autana, Estado Amazonas, Venezuela. Herpetologica, 59, 504–512.

http://dx.doi.org/10.1655/01-31

Barth, R. (1956) Observações anatômicas sôbre a larva de Thoropa miliaris (Amphibia, Leptodactylidae). Memorias do Instituto Oswaldo Cruz, 54, 489–497.

Bionda, C., Gari, N., Luque, E., Salas, N., Lajmanovich, R. & Martino, A. (2012) Trophic ecology in tadpoles of Rhinella arenarum (Anura: Bufonidae) in agroecosystems and their possible implications for conservation. Revista de Biología Tropical, 60, 771–779.
http://dx.doi.org/10.15517/rbt.v60i2.3998

Blackburn, D. (2004) Preliminary analysis of cranial skeletal development in Nectophrynoides vivipara. Journal of Morphology, 260, 279.

http://dx.doi.org/10.1002/jmor.10223

Blackburn, D.C. & Duellman, W.E. (2013) Brazilian marsupial frogs are diphyletic (Anura: Hemiphractidae: Gastrotheca). Molecular Phylogenetics and Evolution, 68, 709–714.

http://dx.doi.org/10.1016/j.ympev.2013.04.021

Blotto, B.L., Nuñez, J.J., Basso, N.G., Úbeda, C.A., Wheeler, W.C. & Faivovich, J. (2013) Phylogenetic relationships of a Patagonian frog radiation, the Alsodes + Eupsophus clade (Anura: Alsodidae), with comments on the supposed paraphyly of Eupsophus. Cladistics, 29, 113–131.

http://dx.doi.org/10.1111/j.1096-0031.2012.00417.x

Bokermann, W.C.A. (1962) Observações biológicas sobre Physalaemus cuvieri Fitz, 1826 (Amphibia, Salientia). Revista Brasileira de Zoologia, 22, 391–399.

Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. 2nd Edition. Taylor and Francis, London.

http://dx.doi.org/10.5962/bhl.title.12075

Boulenger, G.A. (1904) Description of a new tree-frog of the genus Hyla, from British Guiana, carrying eggs on the back. Proceedings of the Zoological Society of London, 1904, 106.

http://dx.doi.org/10.1111/j.1469-7998.1904.tb08315.x

Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795–803.

http://dx.doi.org/10.2307/2408870

Burggren, W.W., Crossley, D.A., Rogowitz, G. & Thompson, D. (2003) Clutch effects explain heart rate variation in embryonic frogs (Cave Coqui, Eleutherodactylus cooki). Physiological and Biochemical Zoology, 76, 672–678.
http://dx.doi.org/10.1086/376918

Brower, A.V.Z. (2006) The how and why of branch support and partitioned branch support, with a new index to assess partition incongruence. Cladistics, 22, 378–386.

http://dx.doi.org/10.1111/j.1096-0031.2006.00113.x

Burton, T.C. (2004) Muscles of the pes of hylid frogs. Journal of Morphology, 260, 209–233.

http://dx.doi.org/10.1002/jmor.10204

Cabrera-Guzmán, E., Crossland, M.R. & Shine, R. (2013) Mechanisms of competition between tadpoles of Australian frogs (Litoria spp.) and invasive cane toads (Rhinella marina). Freshwater Biology, 58, 2584–2600.

http://dx.doi.org/10.1111/fwb.12235

Caldwell, J.P. (2005) A new Amazonian species of Cryptophyllobates (Anura: Dendrobatidae). Herpetologica, 61, 449–461.
http://dx.doi.org/10.1655/04-86.1

Campbell, J.A. (1998) Amphibians and reptiles of northern Guatemala, the Yucatán, and Belize. Animal Natural History Series. Vol. 4. University of Oklahoma Press, Publishing Division of the University, Norman, Oklahoma, 400 pp.

Canedo, C. & Rickli, E. (2006) Female reproductive aspects and seasonality in the reproduction of Eleutherodactylus binotatus (Spix, 1824) (Amphibia, Leptodactylidae) in an Atlantic Rainforest fragment, Southeastern Brazil. Herpetological Review, 37, 149–151.

Caramaschi, U. & Pombal, J.P. (2001) Barycholos savagei: A junior synonym of Paludicola ternetzi, with notes on development. Journal of Herpetology, 35, 357–360.
http://dx.doi.org/10.2307/1566134

Cárdenas-Rojas, D.R., Rabanal, F. & Formas J.R. (2007) The tadpole of Hylorina sylvatica (Anura: Cyclorhamphidae) in southern Chile. Zootaxa, 1580, 51–62.

Cardoso, A.J. & Arzabe, C. (1993) Corte e desenvolvimento larvário de Pleurodema diplolistris (Anura: Leptodactylidae). Revista Brasileira de Biologia, 53, 561–570.

Castañeda, L.E., Sabat, P., Gonzalez, S.P. & Nespolo, R.F. (2006) Digestive plasticity in tadpoles of the Chilean giant frog (Caudiverbera caudiverbera): Factorial effects of diet and temperature. Physiological and Biochemical Zoology, 79, 919–926.
http://dx.doi.org/10.1086/506006

Castro-Herrera, F., Valencia, A. & Villaquirán, D. (2012) Diversidad de anfibios y reptiles del Parque Nacional Natural Isla Gorgona. Feriva Impresores S.A. Cali, Colombia, 112 pp.

Channing, A. (2001) Amphibians of Central and Southern Africa. Cornell University Press, Ithaca, 470 pp.

Channing, A. & Howell, K.M. (2006) Amphibians of East Africa. Edition Chimaira, Frankfurt am Main, 428 pp.

Chibon, P. (1960) Développement au laboratoire d'Eleutherodactylus martinicensis Tschudi, batracien anoure a développement direct. Bulletin de la Société Zoologique de France, 85, 412–418.

Chippindale, P.T., Bonett, R.M., Baldwin, A.S. & Wiens, J.J. (2004) Phylogenetic evidence for a major reversal of life history evolution in plethodontid salamanders. Evolution, 58, 2809–2822.

http://dx.doi.org/10.1111/j.0014-3820.2004.tb01632.x

Cope, E.D. (1866) On the structure and distribution of the genera of the arciferous Anura. Journal of the Academy of Natural Sciences of Philadelphia. Series 2, 6, 67–112.

Crawford, A.J., Cruz, C., Griffith, E., Ross, H., Ibáñez, R., Lips, K.R., Driskell, A.C., Bermingham, E. & Crump, P. (2013) DNA barcoding applied to ex situ tropical amphibian conservation programme reveals cryptic diversity in captive populations. Molecular Ecology Resources, 13, 1005–1018.

http://dx.doi.org/10.1111/1755-0998.12054

Crump, M.L. (1974) Reproductive strategies in a tropical anuran community. Miscellaneous Publications, University of Kansas, Museum of Natural History, 61, 1–68.

http://biodiversitylibrary.org/page/4405402

Darst, C.R. & Cannatella, D.C. (2004) Novel relationships among hyloid frogs inferred from 12S and 16S mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 31, 462–475.

http://dx.doi.org/10.1016/j.ympev.2003.09.003

Davies, M. & Watson, G. (1994) Morphology and reproductive biology of Limnodynastes salmini, L. convexiusculus and Megistolotis lignarius (Anura: Leptodactylidae: Limnodynastinae). Transactions of the Royal Society of South Australia, 118, 149–169.
http://dx.doi.org/10.1670/0022-1511(2002)036[0490:TTOPAA]2.0.CO;2

De la Riva, I. (2007) Bolivian frogs of the genus Phrynopus, with the description of twelve new species (Anura: Brachycephalidae). Herpetological Monographs, 21, 241–277.
http://dx.doi.org/10.1655/07-011.1

De la Riva, I. & Lynch, J.D. (1997) New species of Eleutherodactylus from Boliva (Amphibia: Leptodactylidae). Copeia, 1997, 151–157.
http://dx.doi.org/10.2307/1447850

Del Pino, E.M. (1980) Morphology of the pouch and incubatory integument in marsupial frogs (Hylidae). Copeia, 1980, 10–17.

http://dx.doi.org/10.2307/1444129

Del Pino, E.M. & Escobar, B. (1981) Embryonic stages of Gastrotheca riobambae (Fowler) during maternal incubation and comparison of development with other marsupial frogs. Journal of Morphology, 167, 277–295.

http://dx.doi.org/10.1002/jmor.1051670303

Del Pino, E.M., Venegas-Ferrín, M., Romero-Carvajal, A., Montenegro-Larrea, P., Sáenz-Ponce, N., Moya, I.M., Alarcón, I., Sudou, N., Yamamoto, S. & Taira, M. (2007) A comparative analysis of frog early development. Proceedings of the National Academy of Sciences of the United States of America, 104, 11882–11888.

http://dx.doi.org/10.1073/pnas.0705092104

De Sá, R.O. & Langone, J.A. (2002) The tadpole of Proceratophrys avelinoi (Anura: Leptodactylidae). Journal of Herpetology, 36, 490–494.

http://dx.doi.org/10.1670/0022-1511(2002)036[0490:TTOPAA]2.0.CO;2

Diaz, N.F. & Valencia, J. (1985) Larval morphology and phenetic relationships of the Chilean Alsodes, Telmatobius, Caudiverbera and Insuetophrynus (Anura, Leptodactylidae). Copeia, 1985, 175–181.
http://dx.doi.org/10.2307/1444807

Dixo, M. & Verdade, V.K. (2006) Herpetofauna de serrapilheira da Reserva Florestal de Morro Grande, Cotia (SP). Biota Neotropica, 6, 1–20.
http://dx.doi.org/10.1590/S1676-06032006000200009

Downie, J.R., Livingstone, S.R. & Cormak, J.R. (2001) Selection of the tadpole deposition sites by male Trinidadian stream frogs, Mannophryne trinitatis (Dendrobatidae): an example of antipredator behavior. Herpetological Journal, 11, 91–100.

Dubois, A. (1987 "1986") Miscellanea taxinomica batrachologica (I). Alytes, 5, 7–95.

Duellman, W.E. (1970) The hylid frogs of Middle America. Monograph of the Museum of Natural History, the University of Kansas, 1, 1–753.

http://dx.doi.org/10.5962/bhl.title.2835

Duellman, W.E. (1974) A systematic review of the marsupial frogs (Hylidae: Gastrotheca) of the Andes of Ecuador. Occasional Papers of the Museum of Natural History, the University of Kansas, 22, 1–27.

Duellman, W.E. (1984) Taxonomy of Brazilian hylid frogs of the genus Gastrotheca (Anura: Hylidae). Journal of Herpetology, 18, 302–312.
http://dx.doi.org/10.2307/1564084

Duellman, W.E. (1990) A new species of leptodactylid frog, genus Ischnocnema, from Peru. Occasional Papers of the Museum of Natural History, the University of Kansas, 138, 1–7.

Duellman, W.E. (1997) Amphibians of La Escalera Region, southeastern Venezuela: taxonomy, ecology, and biogeography. Scientific Papers, Natural History Museum, University of Kansas, 2, 1–52.

Duellman, W.E., Barley, A.J. & Venegas, P.J. (2014) Cryptic species diversity in marsupial frogs (Anura: Hemiphractidae: Gastrotheca) in the Andes of northern Peru. Zootaxa, 3768 (2), 159–177.

http://dx.doi.org/10.11646/zootaxa.3768.2.4

Duellman, W.E., Catenazzi, A. & Blackburn, D.C. (2011b) A new species of marsupial frog (Anura: Hemiphractidae: Gastrotheca) from the Andes of southern Peru. Zootaxa, 3095, 1–14.

Duellman, W.E. & De la Riva, I. (1999) Rediscovery and taxonomic status of Hyla splendens Schmidt, 1857 (Anura: Hylidae). Copeia, 1999, 197–199.

http://dx.doi.org/10.2307/1447402

Duellman, W.E. & Fritts, T.H. (1972) A taxonomic review of the southern Andean marsupial frogs (Hylidae: Gastrotheca). Occasional Papers of the Museum of Natural History, the University of Kansas, 9, 1–37.

Duellman, W.E. & Gray, P. (1983) Developmental biology and systematics of the egg-brooding hylid frogs, genera Flectonotus and Fritziana. Herpetologica, 39, 333–359.

http://dx.doi.org/10.2307/3892530

Duellman, W.E. & Hillis, D.M. (1987) Marsupial frogs (Anura: Hylidae: Gastrotheca) of the Ecuadorian Andes: resolution of taxonomic problems and phylogenetic relationships. Herpetologica, 43, 141–173.

http://dx.doi.org/10.2307/3892047

Duellman, W.E. & Hoogmoed, M.S. (1984) The taxonomy and phylogenetic relationships of the hylid frog genus Stefania. Miscellaneous Publications of the Museum of Natural History, University of Kansas, 75, 1–39.

Duellman, W.E., Jungfer, K.-H. & Blackburn, D.C. (2011a) The phylogenetic relationship of geographically separated "Flectonotus" (Anura: Hemiphractidae), as revealed by molecular, behavioral, and morphological data. Phyllomedusa, 10, 15–29.

http://dx.doi.org/10.11606/issn.2316-9079.v10i1p15-29

Duellman, W.E. & Köhler, J. (2005) New species of marsupial frog (Hylidae: Hemiphractinae: Gastrotheca) from the Yungas of Bolivia. Journal of Herpetology, 39, 91–100.

http://dx.doi.org/10.1670/0022-1511(2005)039[0091:NSOMFH]2.0.CO;2

Duellman, W.E. & Lizana, M. (1994) Biology of a sit-and-wait predator, the leptodactylid frog Ceratophrys cornuta. Herpetologica, 51–64.

http://www.jstor.org/stable/3892875

Duellman, W.E. & Lynch, J.D. (1969) Descriptions of Atelopus tadpoles and their relevance to atelopodid classification. Herpetologica, 25, 231–240.

http://www.jstor.org/stable/3891213

Duellman, W.E. & Maness, S.J. (1980) The reproductive behavior of some hylid marsupial frogs. Journal of Herpetology, 14, 213–222.
http://dx.doi.org/10.2307/1563542

Duellman, W.E., Maxson, L.R. & Jesiolowski, C.A. (1988) Evolution of marsupial frogs (Hylidae: Hemiphractinae): Immunological evidence. Copeia, 1988, 527–543.

http://dx.doi.org/10.2307/1445371

Duellman, W.E. & Trueb, L. (1986) Biology of amphibians. McGraw-Hill, New York. 696 pp.

Duellman, W.E. & Venegas, P.J. (2005) Marsupial frogs (Anura: Hylidae: Gastrotheca) from the Andes of northern Peru with descriptions of two new species. Herpetologica, 61, 295–307.

http://dx.doi.org/10.1655/04-105.1

Duméril, A.M.C. & Bibron, G. (1841) Erpétologie genérale ou histoire naturelle complète des reptiles. Vol. 8. Librarie Enclyclopedique de Roret, Paris, 447 pp.

Dunn, E.R. (1937) The amphibian and reptilian fauna of bromeliads in Costa Rica and Panama. Copeia, 1937, 163–167.
http://dx.doi.org/10.2307/1436136

Dunn, E.R. (1944) Notes in Colombian herpetology, II. Caldasia, 2, 397–406.

Elinson, R.P. (2013) Metamorphosis in a frog that does not have a tadpole. In: Shi, Y.-B. (Ed.), Current topics in developmental biology. Academic Press, Burlington, pp. 259–276.
http://dx.doi.org/10.1016/b978-0-12-385979-2.00009-5

Ergueta, P. (1993) Aspectos de la biología y ecología de Phrynopus laplacai (Anura: Leptodactylidae) en un bosque nublado de altura de Yungas (La Paz, Bolivia). Ecología en Bolivia, 21, 19–29.

Faivovich, J. (1998) Comments on the larvae of the Argentine species of the genus Crossodactylus (Leptodactylidae, Hylodinae). Alytes, 16, 61–67.

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.

http://dx.doi.org/10.1206/0003-0090(2005)294[0001:srotff]2.0.co;2

Faivovich, J., Nicoli, L., Blotto, B. L., Pereyra, M.O., Baldo, D., Barrionuevo, J.S., Fabrezi, M., Wild, E.R. & Haddad, C.F. (2014) Big, bad, and beautiful: Phylogenetic relationships of the horned frogs (Anura: Ceratophryidae). South American Journal of Herpetology, 9, 207–227.

http://dx.doi.org/10.2994/SAJH-D-14-00032.1

Farris, J.S. (1983) The logical basis of phylogenetic analysis. In: Platnick, N.I. & Funk, V.A. (Eds), Advances in Cladistics: Proceedings of the Third Meeting of the Willi Hennig Society 2. Columbia University Press, New York, USA, pp. 7–36.

Farris, J.S., Albert, V.A., Källersjö, M., Lipscomb, D. & Kluge, A.G. (1996) Parsimony jackknifing outperforms neighbor-joining. Cladistics, 12, 99–124.

http://dx.doi.org/10.1111/j.1096-0031.1996.tb00196.x

Farris, J.S., Kluge, A.G. & Carpenter, J.M. (2001) Popper and likelihood versus "Popper*". Systematic Biology, 50, 438–444.

http://dx.doi.org/10.1080/10635150119150

Fenolio, D., Hill, R., Kaylock, J. & Cruse, S.J. (2010) The use of a modified shipping container as an amphibian laboratory at the Atlanta Botanical Garden. Leaf Litter, 3, 6–9.

Fisher-Reid, M.C. & Wiens, J.J. (2011) What are the consequences of combining nuclear and mitochondrial data for phylogenetic analysis? Lessons from Plethodon salamanders and 13 other vertebrate clades. BMC Evolutionary Biology, 11, 300.

http://dx.doi.org/10.1186/1471-2148-11-300

Folly, M., Hepp, F., Carvalho-e-Silva, S.P. & Duellman, W.E. (2014) Taxonomic status and redescription of Flectonotus ulei (Anura: Hemiphractidae), with a key for the species of Fritziana. Zoologia (Curitiba), 31, 393–399.

http://dx.doi.org/10.1590/S1984-46702014000400011

Formas, J.R. (1976) Descriptions of Batrachyla (Amphibia, Anura, Leptodactylidae) tadpoles. Journal of Herpetology, 10, 221–225.

http://www.jstor.org/stable/1562983

Formas, J.R., Díaz, N.F. & Valencia, J. (1980) The tadpole of the Chilean frog Insuetophrynus acarpicus. Herpetologica, 36, 316–318.

Formas, J.R. & Pugin, E. (1978) Tadpoles of Eupsophus roseus and Bufo variegatus (Amphibia, Anura) in southern Chile. Journal of Herpetology, 12, 243–246.

http://dx.doi.org/10.2307/1563413

Fouquet, A., Blotto, B.L., Maronna, M.M., Verdade, V.K., Juncá, F A. & de Sá, R. & Rodrigues, M.T. (2013) Unexpected phylogenetic positions of the genera Rupirana and Crossodactylodes reveal insights into the biogeography and reproductive evolution of leptodactylid frogs. Molecular Phylogenetics and Evolution, 67, 445–457.

http://dx.doi.org/10.1016/j.ympev.2013.02.009

Frenkel, C., Vallejo, A., Félix-Novoa, C., Ron, S.R. (2010) Gastrotheca riobambae. In: Ron, S.R., Guayasamin, J.M., Yañez-Muñoz, M.H., Merino-Viteri, A., Ortiz, D.A. (Eds), AmphibiaWebEcuador. Version 2014.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from: http://zoologia.puce.edu.ec/vertebrados/anfibios/FichaEspecie.aspx?Id
=1136 (accesed 12 February 2014)

Frost, D.R. (1985) Amphibian species of the world. A taxonomic and geographical reference. Association of Systematics Collections and Allen Press, Lawrence, 732 pp.

Frost, D.R. (2014) 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 12 October 2014)

Frost, D.R., Grant, T., Faivovich, J., Bain, R., 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. & 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;2

Gaupp, E. (1897) Anatomie des Frosches. Vol. 2. Von Friedrich Vieweg und shon, Braunschweig, 227 pp.

Gerlach, J. (2007) Distribution and status of the Seychelles frogs (Amphibia: Anura: Sooglossidae). Herpetological Journal, 17, 115–122.

Giaretta, A.A. (1996) Reproductive specializations of the bromeliad hylid frog Phyllodytes luteolus. Journal of Herpetology, 30, 96–97.

http://dx.doi.org/10.2307/1564719

Giaretta, A.A. & Facure, K.G. (2008) Reproduction and habitat of ten Brazilian frogs (Anura). Contemporary Herpetology, 2008, 1–4.

Giaretta, A.A & Gomes, F.K. (2004) Reproductive ecology and behavior of Thoropa miliaris (Spix, 1824) (Anura, Leptodactylidae, Telmatobiinae). Biota Neotropica, 4, 1–10.
http://dx.doi.org/10.1590/S1676-06032004000200008

Gitlin, D. (1944) The development of Eleutherodactylus portoricensis. Copeia, 1944, 91–98.

http://dx.doi.org/10.2307/1438760

Glaw, F. & Franzen, M. (2006) Type catalogue of amphibians in the Zoologische Staatsammlung München. Spixiana, 29, 153–192.

Goin, C.J. (1947) Studies on the life history of Eleutherodactylus ricordii planirostris (Cope) in Florida: with special reference to the local distribution of an allelomorphic color pattern. University of Florida Press, Gainesville, Florida, 66 pp

Goloboff, P.A. (1999) Analyzing large data sets in reasonable times: solutions for composite optima. Cladistics, 15, 415–428.

http://dx.doi.org/10.1111/j.1096-0031.1999.tb00278.x

Goloboff, P.A., Carpenter, J.M., Arias, J.S. & Esquivel, D.R.M. (2008b) Weighting against homoplasy improves phylogenetic analysis of morphological data sets. Cladistics, 24, 758–773.

http://dx.doi.org/10.1111/j.1096-0031.2008.00209.x

Goloboff, P.A., Farris, J.S. & Nixon, K.C. (2008a) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774–786.

http://dx.doi.org/10.1111/j.1096-0031.2008.00217.x

Gomez-Mestre, I., Pyron, R.A. & Wiens, J.J. (2012) Phylogenetic analyses reveal unexpected patterns in the evolution of reproductive modes in frogs. Evolution, 66, 3687–3700.

http://dx.doi.org/10.1111/j.1558-5646.2012.01715.x

González, E., Galvani, G., Sanabria, E., Barrasso, D.A., Alcalde, L. & Quiroga, L. (2014) The tadpole of Odontophrynus barrioi Cei, Ruiz, and Beçak, 1982 (Anura: Odontophrynidae): a comparison with the other tadpoles of the genus. Acta Herpetologica, 9, 15–23.

http://dx.doi.org/10.13128/Acta_Herpetol-12931

Goodman, M., Olson, C.B., Beeber, J.E. & Czelusniak, J. (1982) New perspectives in the molecular biological analysis of mammalian phylogeny. Acta Zoologica Fennica, 169, 19–35.

Gosner, K.L., 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica, 16, 183–190.
http://dx.doi.org/10.1206/0003-0090(2006)299[1:PSODFA]2.0.CO;2

Graham, A. (2009) The Andes: a geological overview from a biological perspective. Annals of the Missouri Botanical Garden, 96, 371–385.

http://dx.doi.org/10.3417/2007146

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

http://dx.doi.org/10.1206/0003-0090(2006)299[1:psodfa]2.0.co;2

Grant, T. & Kluge, A.G. (2004) Transformation series as an ideographic character concept. Cladistics, 20, 23–31.

http://dx.doi.org/10.1111/j.1096-0031.2004.00003.x

Grant, T. & Kluge, A.G. (2007) Ratio of explanatory power (REP): a new measure of group support. Molecular Phylogenetics and Evolution, 44, 483–487.

http://dx.doi.org/10.1016/j.ympev.2006.11.027

Grant, T. & Kluge, A.G. (2008a) Credit where credit is due: the Goodman-Bremer support metric. Molecular Phylogenetics and Evolution, 49, 405–406.

http://dx.doi.org/10.1016/j.ympev.2008.04.023

Grant, T. & Kluge, A.G. (2008b) Clade support measures and their adequacy. Cladistics, 24, 1051–1064.

http://dx.doi.org/10.1111/j.1096-0031.2008.00231.x

Grant, T. & Kluge, A.G. (2009) Perspective: parsimony, explanatory power, and dynamic homology testing. Systematics and Biodiversity, 7, 357–363.

http://dx.doi.org/10.1017/S147720000999017X

Gravenhorst, J.L.C. (1825) Stombus, eine neue Amphibien Gattung. Isis von Oken, 1825, 920–922.

Guayasamin, J.M., Castroviejo-Fisher, S., Ayarzagüena, J., Trueb, L. & Vilà, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574–595.

http://dx.doi.org/10.1016/j.ympev.2008.04.012

Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzagüena, J., Rada, M. & Vilà, C. (2009) Phylogenetic systematics of glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1–97.

http://dx.doi.org/10.1016/j.ympev.2008.04.012

Günther, A.C.L.G. (1859 "1858") Catalogue of the Batrachia Salientia in the Collection of the British Museum. Taylor and Francis, London, 160 pp.

Haas, A. (2003) Phylogeny of frogs as inferred from primarily larval characters (Amphibia: Anura). Cladistics, 19, 23–90.

http://dx.doi.org/10.1016/s0748-3007(03)00006-9

Haad, B., Candioti, F.V. & Baldo, D. (2011) Shape variation in lentic and lotic tadpoles of Melanophryniscus (Anura: Bufonidae). Studies on Neotropical Fauna and Environment, 46, 91–99.

http://dx.doi.org/10.1080/01650521.2011.593124

Haddad, C.F.B. & Prado, C.P.A. (2005) Reproductive modes in frogs and their unexpected diversity in the Atlantic Forest of Brazil. BioScience, 55, 207–217.

http://dx.doi.org/10.1641/0006-3568(2005)055[0207:RMIFAT]2.0.CO;2

Hall, B.K. (2000) Balfour, Garstang and de Beer: the first century of evolutionary embryology. American Zoologist, 40, 718–728.

http://dx.doi.org/10.1093/icb/40.5.718

Hanken, J. (2003) Direct develpment. In: Hall, B.K. & Olson, W.M. (Eds.), Keywords and concepts in evolutionary developmental biology. Harvard University Press, Cambridge, pp. 97–102.

Harper, E.B., Measey, G.J., Patrick, D.A., Menegon, M. & Vonesh, J.R. (2010) Field guide to the amphibians of the Eastern Arc Mountains and Coastal Forests of Tanzania and Kenya. Camerapix, Nairobi. 143 pp.

Hartmann, M.T., Hartmann, P.A. & Haddad, C.F.B. (2010) Reproductive modes and fecundity of an assemblage of anuran amphibians in the Atlantic rainforest, Brazil. Iheringia, Série Zoologia, 100, 207–215.
http://dx.doi.org/10.1590/S0073-47212010000300004

Heatwole, H. (1962) Contributions to the natural history of Eleutherodactylus cornutus maussi. Stahlia Miscellaneous Papers, 2, 1–11.

Hedges, S.B. (1990) A new species of Phrynopus (Anura: Leptodactylidae) from Peru. Copeia, 1990, 108–112.
http://dx.doi.org/10.2307/1445826

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., 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.

Heyer, W.R. (1969) Studies on the genus Leptodactylus (Amphibia, Leptodactylidae) III. A redefinition of the genus Leptodactylus and a description of a new genus of leptodactylid frogs. Contributions in Science. Natural History Museum of Los Angeles County, 155, 1–14.

Hill, R., Kaylock, J., Griffith, E., Ross, H., Gagliardo, R., Timpe, E. & Crump, P. (2010) Observations on the captive reproduction of Gaige’s Rain Frog, Pristimantis gaigeae. Herpetological Review, 41, 465–467.

Hödl, W. (1990) Reproductive diversity in Amazonian lowland frogs. Fortschritte der Zoologie, 38, 41–60.

Hoffmann, H. (2006) Observations on behaviour and parental care of Leptodactylus melanonotus (Hallowell) in Costa Rica. Salamandra, 42, 109–116.

Hoffman, H. (2010) The glass frog tadpoles of Costa Rica (Anura: Centrolenidae): a study of morphology. Abhandlungen der Senckenberg Gesellschaft für Naturforschung, 567, 1–78.

Hoogmoed, M.S. & Gruber, U. (1983) Spix and Wagler type specimens of reptiles and amphibians in the Natural History Musea in Munich (Germany) and Leiden (the Netherlands). Spixiana, 9, 319–415.

Hovenkamp, P. (2004) In support of support. In: Stevenson, D.W. (Ed.), Abstracts of the 22nd Annual Meeting of the Will Hennig Society. Cladistics, 20, 76–100.

http://dx.doi.org/10.1111/j.1096-0031.2003.00006.x

Hovenkamp, P. (2009) Support and stability. Cladistics, 25, 107–108.

http://dx.doi.org/10.1111/j.1096-0031.2008.00243.x

Hsu, M.-Y., Kam, Y.-C. & Fellers, G.M. (2006) Temporal organization of an anuran acoustic community in a Taiwanese subtropical forest. Journal of Zoology, 269, 331–339.

http://dx.doi.org/10.1111/j.1469-7998.2006.00044.x

Hughes, A. (1959) Studies in embryonic and larval development in Amphibia I. The embryology of Eleutherodactylus ricordii, with special reference to the spinal cord. Journal of Embryology and Experimental Morphology, 7, 22–38.

Infante-Rivero, E.E., Rojas-Runjaic, F.J.M. & Barrio-Amorós, C.L. (2009 “2008”) Un nuevo Cryptobatrachus Ruthven, 1916 (Anura, Cryptobatrachidae) de la vertiente venezolana de la Sierra de Perijá. Memoria de la Fundación La Salle de Ciencias Naturales, 169, 45–63.

Izecksohn, E., Carvalho-e-Silva, S.P. de & Peixoto, O.L. (2009) Sobre Gastrotheca fissipes (Boulenger, 1888), com descrição de uma nova espécie (Amphibia, Anura, Amphignathodontidae). Arquivos do Museu Nacional, Rio de Janeiro, 67, 81–91.

Iwai, N. & Kagaya, T. (2005) Growth of Japanese Toad (Bufo japonicus formosus) tadpoles fed different food items. Current Herpetology, 24, 85–89.

http://dx.doi.org/10.3105/1345-5834(2005)24[85:GOJTBJ]2.0.CO;2

Jameson, D.L. (1950) The development of Eleutherodactylus latrans. Copeia, 1950, 44–46.
http://dx.doi.org/10.2307/1437581

Jameson, D.L. (1955) The population dynamics of the Cliff Frog, Syrrhophus marnocki. American Midland Naturalist, 54, 342–381.
http://dx.doi.org/10.2307/2422572

Joglar, R.L., Burrowes, P.A. & Rios, N. (1996) Biology of the Puerto Rican cave-dwelling frog, Eleutherodactylus cooki, and some recommendations for its conservation. In: Powell, R. & Henderson, R.W. (Eds.), Contributions to West Indian Herpetology: a Tribute to Albert Schwartz. Society for the Study of Amphibians and Reptiles, Contributions to Herpetology. Vol. 12. Ithaca, New York, pp. 251–258.

Johnson, L.M. (1991) Growth and development of larval northern cricket frogs (Acris crepitans) in relation to phytoplankton abundance. Freshwater Biology, 25, 51–59.

http://dx.doi.org/10.1111/j.1365-2427.1991.tb00472.x

Jorquera, B., Garrido, O., & Pugin, E. (1982) Comparative studies of the digestive tract development between Rhinoderma darwinii and R. rufum. Journal of Herpetology, 16, 204–214.

http://dx.doi.org/10.2307/1563714

Kaefer, I.L., Both, C. & Cechin, S.Z. (2009) Breeding biology of the rapids frog Limnomedusa macroglossa (Anura: Cycloramphidae) in southern Brazil. Journal of Natural History, 43, 1195–1206.

http://dx.doi.org/10.1080/00222930902767474

Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution, 30, 772–780.

http://dx.doi.org/10.1093/molbev/mst010

Kenny, J. S. (1969) The Amphibia of Trinidad. Studies on the Fauna of Curaçao and Other Caribbean Islands, 29, 1–78.

Kerney, R.R., Blackburn, D.C., Müller, H. & Hanken, J. (2012) Do larval traits re-evolve? Evidence from the embryogenesis of a direct-developing salamander, Plethodon cinereus. Evolution, 66, 252–262.

http://dx.doi.org/10.1111/j.1558-5646.2011.01426.x.

Kerney, R., Meegaskumbura, M., Manamendra-Arachchi, K. & Hanken, J. (2007) Cranial ontogeny in Philautus silus (Anura: Ranidae: Rhacophorinae) reveals few similarities with other direct-developing anurans. Journal of Morphology, 268, 715–725.

http://dx.doi.org/10.1002/jmor.10545

Kluge, A.G. (1989) A concern for evidence and a phylogenetic hypothesis of relationships among Epicrates (Boidae, Serpentes). Systematic Biology, 38, 7–25.

http://dx.doi.org/10.1093/sysbio/38.1.7

Kluge, A.G. (2001a) Philosophical conjectures and their refutation. Systematic Biology, 50, 322–330.

http://dx.doi.org/10.1080/10635150119615

Kluge, A.G. (2001b) Parsimony with and without scientific justification. Cladistics, 17, 199–210.

http://dx.doi.org/10.1111/j.1096-0031.2001.tb00117.x

Kluge, A.G. & Grant, T. (2006) From conviction to anti-superfluity: old and new justifications for parsimony in phylogenetic inference. Cladistics, 22, 276–288.

http://dx.doi.org/10.1111/j.1096-0031.2006.00100.x

Köhler, J. (2000) Amphibian diversity in Bolivia: a study with special reference to montane forest regions. Bonner Zoologische Monographien, 48, 1–243.

Kok, P.J.R. & Kalamandeen, M. (2008) Introduction to the taxonomy of the amphibians of Kaieteur National Park, Guyana. Abc Taxa, 5, 1–278.

Kok, P.J.R., MacCulloch, R.D., Means, D.B., Roelants, K., van Bocxlaer, I. & Bossuyt, F. (2012) Low genetic diversity in tepui summit vertebrates. Curreent Biology, 22, 1–25.

http://dx.doi.org/10.1016/j.cub.2012.06.034

Konstant, B. (2007) News from the field safeguarding Panama’s amphibians. Connect, 2007, 22.

Kurth, M., Hörnes, D. & Rödder, D. (2013) Race against desiccation: rapid larval development in Melanophryniscus klappenbachi (Anura: Bufonidae). Salamandra, 50, 117–124.

Kwet, A., Lingnau, R. & Di-Bernardo, M. (2010) Pró-Mata: Anfíbios da Serra Gaúcha, Sul do Brasil. Universität Tübingen, Tübingen, 148 pp.

Lamar, W.W. & Wild, E.R. (1995) Comments on the natural history of Lithodytes lineatus (Anura: Leptodactylidae), with description of the tadpole. Herpetological Natural History, 3, 135–142.

Langone, J.A. (1989) Descripción de la larva de Physalaemus gracilis (Boulenger, 1883) (Amphibia, Anura, Leptodactylidae). Comunicaciones Zoológicas del Museo de Historia Natural de Montevideo, 171, 1–12.

Laufer, G. & Barreneche, J.M. (2008) Re-description of the tadpole of Pseudopaludicola falcipes (Anura: Leiuperidae), with comments on larval diversity of the genus. Zootaxa, 1760, 50–58.

Lavilla, E.O. (1988) Lower telmatobiinae (Anura: Leptodactylidae): generic diagnoses based on larval characters. Occasional Papers of the Museum of Natural History, the University of Kansas, 124, 1–19.

Lavilla, E.O. (1990) The tadpole of Hyla nana. Journal of Herpetology, 24, 207–209.

http://dx.doi.org/10.2307/1564233

Lisboa, B.S., Nascimento, F.A.C. do & Skuk, G.O. (2011) Redescription of the tadpole of Macrogenioglottus alipioi (Anura: Cycloramphidae), a rare and endemic species of the Brazilian Atlantic Forest. Zootaxa, 3046, 67–68.

Lutz, B. (1947) Trends towards non-aquatic and direct development in frogs. Copeia, 1947, 242–252.

http://www.jstor.org/stable/1438920

Lutz, B. (1958) Anfíbios novos e raros das serras costeiras do Brasil (New or rare frogs from the coastal ranges of Brazil. Eleutherodactylus venancioi n.sp., E. hoehnei n.sp. Holoaden bradei n.sp. e H. lüderwaldti Mir. Rib. Memórias do Instituto Oswaldo Cruz, 56, 373–405.

Lutz, B. (1968) Taxonomy of the Neotropical Hylidae. Pearce-Sellards Series, Texas Memorial Museum, 11, 3–26.

Lynch, J.D. (1975) A review of the Andean leptodactylid frog genus Phrynopus. Occasional Papers of the Museum of Natural History, the University of Kansas, 35, 1–51.

Lynch, J.D. (1976) Two new species of frogs of the genus Euparkerella (Amphibia: Leptodactylidae) from Ecuador and Peru. Herpetologica, 32, 48–53.

Lynch, J.D. (1981) Leptodactylid frogs of the genus Eleutherodactylus in the Andes of northern Ecuador and adjacent Colombia. Miscellaneous Publication. Museum of Natural History, the University of Kansas, 72, 1–46.

Lynch, J.D. (2006) The tadpoles of frogs and toads found in the lowlands of northern Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 30, 443–457.

Lynch, J.D. (2008) A taxonomic revision of frogs of the genus Cryptobatrachus (Anura: Hemiphractidae). Zootaxa, 1883, 28–68.

Lynch, J.D. & Suárez M., A.M. (2011) Clave ilustrada de los renacuajos en las tierras bajas al oriente de los Andes, con énfasis en Hylidae. Caldasia, 33, 235–270.

Lynn, W.G. (1942) The embryology of Eleutherodactylus nubicola, an anuran which has no tadpole stage. Contributions to Embryology. Carnegie Museum, 190, 29–62.

Lynn, W.G. & Lutz, B. (1946) The development of Eleutherodactylus guentheri Steindachner 1864 (Salientia). Boletim do Museu Nacional Brasil, New Series, 71, 1–46.

Martinazzo, L.B., Basso, N.G. & Úbeda, C.A. (2011) The aquatic and littoral forms of the Patagonian frog Atelognathus patagonicus (Batrachylinae): new molecular evidence. Zootaxa, 3129, 62–68.

Martínez R., C.C., Bell, J. & Parker, J. (2011) Haiti’s disappearing frogs. Amphibian Ark Newsletter, 16, 15–16.

Martins, I.A. (2010) Natural history of Holoaden luederwaldti (Amphibia: Strabomantidae: Holoadeninae) in southeastern of Brazil. Zoologia, 27, 40–46.
http://dx.doi.org/10.1590/S1984-46702010000100007

MacCulloch, R.D. & Lathrop, A. (2002) Exceptional diversity of Stefania (Anura: Hylidae) on Mount Ayanganna, Guyana: three new species and new distribution records. Herpetologica, 58, 327–346.
http://dx.doi.org/10.1655/0018-0831(2002)058[0327:EDOSAH]2.0.CO;2

MacCulloch, R.D. & Amy Lathrop, A. (2005) Hylid frogs from Mount Ayanganna, Guyana: new species, redescriptions, and distributional records. Phyllomedusa, 4, 17–37.

http://dx.doi.org/10.11606/issn.2316-9079.v4i1p17-37

McDiarmid, R.W. & Altig, R. (1999) Tadpoles: the biology of anuran larvae. University of Chicago Press, Chicago, 458 pp.

Melchiors, J., Di-Bernardo, M., Funk P., G.M., Baptista de O., R., Sole, M. & Kwet, A. (2004) Reprodução de Pseudis minuta (Anura, Hylidae) no sul do Brasil. Phyllomedusa, 3, 61–68.
http://dx.doi.org/10.11606/issn.2316-9079.v3i1p61-68

Mendelson, J.R., III, Silva, H.R. da & Maglia, A. M. (2000) Phylogenetic relationships among marsupial frog genera (Anura: Hylidae: Hemiphractinae) based on evidence from morphology and natural history. Zoological Journal of the Linnean Society, 128, 125–148.

http://dx.doi.org/10.1111/j.1096-3642.2000.tb00159.x

Miranda-Ribeiro, A. (1920) As Hyla coelonotas do Museu Paulista. Revista do Museu Paulista, 12, 323–328.

Miranda-Ribeiro, A. (1926) Notas para servirem ao estudo dos Gymnobatrachios (Anura) Brasileiros. Arquivos do Museu Nacional. Rio de Janeiro, 27, 1–227.

Mitchell, N.J. & Seymour, R.S. (2000) Effects of temperature on the energy cost and timing of embryonic and larval development of the terrestrially breeding moss frog, Bryobatrachus nimbus. Physiological and Biochemical Zoology, 73, 829–840.
http://dx.doi.org/10.1086/318097

Mogali, S.M., Saidapur, S.K. & Shanbhag, B.A. (2011) Levels of predation modulate antipredator defense behavior and metamorphic traits in the toad Bufo melanostictus. Journal of Herpetology, 45, 428–431.

http://dx.doi.org/10.1670/10-273.1

Molina R., C.R. (2004 "2002") Reproducción de Pleurodema brachyops (Anura: Lectodactylidae) en los llanos del estado Apure, Venezuela. Memoria de la Fundación La Salle Ciencias Naturales, 158, 117–125.

Montealegre-Delgado, X.K., Avendaño-Casadiego, K. & Bernal, M.H. (2013) Efecto del tamaño del hábitat en la supervivencia, desarrollo y crecimiento en renacuajos de Engystomops pustulosus (Anura: Leiuperidae) y Rhinella humboldti (Anura: Bufonidae). Papéis Avulsos de Zoologia, 53, 209–314.

http://dx.doi.org/10.1590/S0031-10492013002300001

Mora, A., Baby, P., Roddaz, M., Parra, M., Brusset, S., Hermoza, W. & Espurt, N. (2010) Tectonic history of the Andes and sub-Andean zones: implications for the development of the Amazon drainage basin. In: Hoorn, C. & Wesselingh F. (Eds.), Amazonia, landscape and species evolution: a look into the past. Wiley-Blackwell, Chichester, pp. 38–60.

Müller, H. (2006) Ontogeny of the skull, lower jaw, and hyobranchial skeleton of Hypogeophis rostratus (Amphibia: Gymnophiona: Caeciliidae) revisited. Journal of Morphology, 267, 968–986.

http://dx.doi.org/10.1002/jmor.10454

Müller, H., Loader, S.P., Ngalason, W., Howell, K.M. & Gower, D.J. (2007) Reproduction in brevicipitid frogs (Amphibia: Anura: Brevicipitidae)-Evidence from Probreviceps m. macrodactylus. Copeia, 2007, 726–733.
http://dx.doi.org/10.1643/0045-8511(2007)2007[726:RIBFAA]2.0.CO;2

Myers, C.W. (1969) The ecological geography of cloud forest in Panama. American Museum Novitates, 2396, 1–52.

Myers, G.S. & Carvalho, A.L. de (1945) Notes on some new or little-known Brazilian amphibians, with an examination of the history of the Plata salamander, Ensatina platensis. Boletim do Museu Nacional, 35, 1–24.

Natale, G.S., Alcalde, L., Herrera, R., Cajade, R., Schaefer, E.F., Marangoni, F. & Trudeau, V.L. (2010) Underwater acoustic communication in the macrophagic carnivorous larvae of Ceratophrys ornata (Anura: Ceratophryidae). Acta Zoologica, 92, 46–53.

http://dx.doi.org/10.1111/j.1463-6395.2009.00445.x

Nixon, K.C. (1999) The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics, 15, 407–414.

http://dx.doi.org/10.1111/j.1096-0031.1999.tb00277.x

Noble, G.K. (1927) The value of life history data in the study of the evolution of the Amphibia. Annals of the New York Academy of Sciences, 30, 31–128.

http://dx.doi.org/10.1111/j.1749-6632.1927.tb55359.x

Noble, G.K. (1931) The biology of the Amphibia. McGraw-Hill, New York, 577 pp.

Nokhbatolfoghahai, M. & Downie, J.R. (2008) The external gills of anuran amphibians: comparative morphology and ultrastructure. Journal of Morphology, 269, 1197–1213.

http://dx.doi.org/10.1002/jmor.10655

Nokhbatolfoghahai, M., Mitchell, N.J. & Downie, J.R. (2010) Surface ciliation and tail structure in direct-developing frog embryos: a comparison between Myobatrachus gouldii and Pristimantis (= Eleutherodactylus) urichi. Herpetological Journal, 20, 59–68.

Nuin, P.A.S. (2003) Description of the tadpole of Megaelosia goeldii (Leptodactylidae, Hylodinae) with natural history notes. Herpetological Review, 34, 27–28.

Olson, C.A., Beard, K.H. & Pitt, W.C. (2012) Biology and impacts of Pacific island invasive species. 8. Eleutherodactylus planirostris, the Greenhouse Frog (Anura: Eleutherodactylidae). Pacific Science, 66, 255–270.
http://dx.doi.org/10.2984/66.3.1

Ovaska, K. & Rand, A.S. (2001) Courtship and reproductive behavior of the frog Eleutherodactylus diastema (Anura: Leptodactylidae) in Gamboa, Panama. Journal of Herpetology, 35, 44–50.
http://dx.doi.org/10.2307/1566021

Padial, J.M., Chaparro, J.C., Castroviejo-Fisher, S., Guayasamin, J.M., Lehr, E., Delgado C., A.J., Vaira, M., Teixeira Jr., M., Aguayo-Vedia, C.R. & De la Riva, I. (2012) A revision of species diversity in the Neotropical genus Oreobates (Anura: Strabomantidae), with the description of three new species from the Amazonian slopes of the Andes. American Museum Novitates, 3752, 1–55.
http://dx.doi.org/10.1206/3752.2

Padial, 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.3825.1.1

Peracca, M.G. (1914) Reptiles et batraciens de Colombie. Memoires de la Société Neuchateloise de Sciences Naturelles, 5, 96–111.

Peters, W.C.H. (1862) Über die Batrachier-Gattung Hemiphractus. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1862, 144–152.

Peters, W.C.H. (1876) Über eine von Herrn Viceconsul L. Krug und Dr. J. Gundlach auf der Insel Puertorico gemachte Sammlung von Säugethieren und Amphibien, so eire über die Entwickelung eines Batrachiers, Hylodesmartinicensis Dum. Bibr., ohne Metamorphose. Monatsberichte der Königlichen Preussische Akademie des Wissenschaften zu Berlin, 1876, 703–714, 914.

Pombal, J.P. Jr. (1999) Oviposição e desenvolvimento de Brachycephalus ephippium (Spix) (Anura, Brachycephalidae). Revista Brasileira de Zoologia, 16, 967–976.

Pombal, J.P. Jr. & Gasparini, J.L. (2006) A new Brachycephalus (Anura: Brachycephalidae) from the Atlantic Rainforest of Espirito Santo, Southeastern Brazil. South American Journal of Herpetology, 1, 87–93.
http://dx.doi.org/10.1590/S0101-81751999000400004

Pombal, J.P. Jr. & Haddad, C.F.B. (1999) Frogs of the genus Paratelmatobius (Anura: Leptodactylidae) with descriptions of two new species. Copeia, 1999, 1014–1026.

http://dx.doi.org/10.2307/1447976

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, 543–583.

http://dx.doi.org/10.1016/j.ympev.2011.06.012

Quinzio, S.I., Fabrezi, M. & Faivovich, J. (2006) Redescription of the tadpole of Chacophrys pierottii (Vellard, 1948) (Anura, Ceratophryidae). South American Journal of Herpetology, 1, 202–209.

http://dx.doi.org/10.2994/1808-9798(2006)1[202:ROTTOC]2.0.CO;2

Rivero, J.A. (1970) On the origin, endemism and distribution of the genus Stefania Rivero (Amphibia, Salientia) with a description of a new species from southeastern Venezuela. Boletín de la Sociedad Venezolana de Ciencias Naturales, 28, 456–481.

Roelants, K., Gower, D.J., Wilkinson, M., Loader, S.P., Biju, S.D., Guillaume, K., Moriau, L. & Bossuyt, F. (2007) Global patterns of diversification in the history of modern amphibians. Proceedings of the National Academy of Sciences of the United States of America, 104, 887–892.

http://dx.doi.org/10.1073/pnas.0608378104

Ron, S.R. & Frenkel, C. (2010) Gastrotheca pseustes. In: Ron, S.R., Guayasamin, J.M., Yanez-Muñoz, M.H., Merino-Viteri, A. & Ortiz, D.A. (Eds.), AmphibiaWebEcuador. Version 2014.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador. Available from: http://zoologia.puce.edu.ec/vertebrados/anfibios/FichaEspecie.aspx?Id=1134 ( accesed 12 February 2014)

Rossa-Feres, D.C. & Nomura, F. (2006) Characterization and taxonomic key for tadpoles (Amphibia: Anura) from the northwestern region of São Paulo State, Brazil. Biota Neotropica, 6, 1–6.
http://dx.doi.org/10.1590/S1676-06032006000100014

Roth, A.H. & Jackson, J.F. (1987) The effect of pool size on recruitment of predatory insects and on mortality in a larval anuran. Herpetologica, 43, 224–232.

http://www.jstor.org/stable/3892055

Ruibal, R. & Thomas, E. (1988) The obligate carnivorous larvae of the frog, Lepidobatrachus laevis (Leptodactylidae). Copeia, 1988, 591–604.
http://www.jstor.org/stable/1445377

Ruiz-Carranza, P.M., Ardila-Robayo, M.C., Lynch, J.D. & Restrepo-Toro, J.H. (1997) Una nueva especie de Gastrotheca (Amphibia: Anura: Hylidae) de la Cordillera Occidental de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 21, 373–378.

Ruthven, A.G. (1916) A new genus and species of amphibian of the family Cystignathidae. Occasional Papers of the Museum of Zoology, University of Michigan, 33, 1–4.

http://dx.doi.org/2027.42/56472

Ruvinsky, I. & Maxson, L.R. (1996) Phylogenetic relationships among bufonoid frogs (Anura: Neobatrachia) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 5, 533–547.

http://dx.doi.org/10.1006/mpev.1996.0048

Sampson, L.V. (1904) A contribution to the embryology of Hylodes martinicensis. American Journal of Anatomy, 3, 473–504.

http://dx.doi.org/10.1002/aja.1000030404

Sankoff, D.M. (1975) Minimal mutation trees of sequences. SIAM Journal of Applied Mathematics, 28, 35–42.

http://dx.doi.org/10.1137/0128004

Sankoff, D.M. & Rousseau, P. (1975) Locating the vertices of a Steiner tree in arbitrary space. Mathematical Programming, 9, 240–246.

http://dx.doi.org/10.1007/bf01681346

Sankoff, D., Cedergren, R.J. & Lapalme, G. (1976) Frequency of insertion-deletion, transversion, and transition in the evolution of 5s ribosomal rna. Journal of Molecular Evolution, 7, 133–149.

http://dx.doi.org/10.1007/bf01732471

San Mauro, D., Gower, D.J., Müller, H., Loader, S.P., Zardoya, R., Nussbaum, R.A. & Wilkinson, M. (2014) Life-history evolution and mitogenomic phylogeny of caecilian amphibians. Molecular Phylogenetics and Evolution, 73, 177–189.

http://dx.doi.org/10.1016/j.ympev.2014.01.009

Savage, J.M. (2002) The Amphibians and Reptiles of Costa Rica. A herpetofauna between two continents, between two seas. University of Chicago Press, Chicago & London, 943 pp.

Schlaepfer, M.A. & Figueroa-Sandy, R. (1998) Female reciprocal calling in a Costa Rican leaf-litter frog, Eleutherodactylus podiciferus. Copeia, 1998, 1076–1080.
http://dx.doi.org/10.2307/1447362

Schmid, M., Steinlein, C., Bogart, J.P., Feichtinger, W., Haaf, T., Nanda, I., del Pino, E. M., Duellman, W.E. & Hedges, S.B. (2013 "2012") The hemiphractid frogs. Phylogeny, embryology, life history, and cytogenetics. Cytogenetic and Genome Research, 13, 69–384.

http://dx.doi.org/10.1159/000343460

Schlüter, A. (1990) Reproduction and tadpole of Eladorhina perezi (Amphibia, Leptodactylidae). Studies on Neotropical Fauna and Environment, 25, 49–56.
http://dx.doi.org/10.1080/01650529009360801

Señaris, J.C., Ayarzagüena, J. & Gorzula, S.J. (1997 "1996") Revisión taxonómica del genero Stefania (Anura: Hylidae) en Venezuela con la descripción de cinco nuevas especies. Publicaciones de la Asociación Amigos de Doñana, 7, 1–57.

Sheil, C.A. & Mendelson III, J.R. (2001) A new species of Hemiphractus (Anura: Hylidae: Hemiphractinae), and a redescription of H. johnsoni. Herpetologica, 57, 189–202.

Sheil, C.A., Mendelson, J.R. III & Silva H.R. da (2001) Phylogenetic relationships of the species of Neotropical horned frogs, genus Hemiphractus (Anura: Hylidae: Hemiphractinae), based on evidence from morphology. Herpetologica, 57, 203–214.

Siddall, M. (2002) Measures of support. In: DeSalle, R., Giribet, G., Wheeler, W. (Eds.), Techniques in Molecular Systematics and Evolution. Birkhäuser-Verlag, Basel, Switzerland, pp. 81–101.
http://dx.doi.org/10.1007/978-3-0348-8125-8_5

Silva H.R. da (1998) Phylogenetic relationships of the family Hylidae with emphasis on the relationships within the subfamily Hylinae (Amphibia: Anura). Ph.D. dissertation, Department of Systematics and Ecology, University of Kansas. [unkown pagination]

Sinsch, U. & Juraske, N. (2006) Advertisement calls of hemiphractine marsupial frogs: IV. Stefania spp. Proceedings of the 13th Ordinary General Meeting of the Societas Europaea Herpetologica, Bonn, 159–162.

Smith, W.L. & Wheeler, W.C. (2006) Venom evolution widespread in fishes: a phylogenetic road map for the bioprospecting of piscine venoms. Journal of Heredity, 97, 206–217.

http://dx.doi.org/10.1093/jhered/esj034

Spraguea, J. & Zimkus, B.M. (2011) Description of the tadpole of Amietophrynus brauni (Anura: Bufonidae) from the west Usambara mountains, Tanzania. African Journal of Herpetology, 60, 70–77.

http://dx.doi.org/10.1080/21564574.2010.542776

Stuart, S.N., Hoffmann, M., Chanson, J.S., Cox, N.A., Berridge, R.J., Ramani, P. & Young, B.E. (Eds.) (2008) Threatened amphibians of the world. Lynx Edicions, Barcelona, Spain; IUCN, Gland, Switzerland; and Conservation International, Arlington, Virginia, 758 pp.

Sunyer, J., Páiz, G., Matthias Dehling, D. & Köhler, G. (2009) A collection of amphibians from Río San Juan, southeastern Nicaragua. Herpetology Notes, 2, 189–202.

Teixeira Jr., M., Vechio, F.D., Recoder, R.S., Carnaval, A.C., Strangas, M., Damasceno, R.P., Sena, M.A. de & Rodrigues, M.T. (2012) Two new species of marsupial tree-frogs genus Gastrotheca Fitzinger, 1843 (Anura, Hemiphractidae) from the Brazilian Atlantic Forest. Zootaxa, 3437, 1–23.

Thibaudeau, G. & Altig, R. (1999) Endotrophic anurans: development and evolution. In: McDiarmid, R.W. & Altig, R. (Eds), Tadpoles: The biology of anuran larvae. University of Chicago Press, Chicago, pp. 170–188.

Torres-Orozco, R.E., Jimenez-Sierra, C.L., Vogt, R.C., Villarreal-Benitez, J.L. (2002) Neotropical tadpoles: spatial and temporal distribution and habitat use in a seasonal lake in Veracruz, Mexico. Phyllomedusa, 1, 81–91.
http://dx.doi.org/10.11606/issn.2316-9079.v1i2p81-91

Townsend, D.S. & Stewart, M.M. (1985) Direct development in Eleutherodactylus coqui (Anura: Leptodactylidae): a staging table. Copeia, 1985, 423–436.

http://www.jstor.org/stable/1444854

Trueb, L. (1970) Evolutionary relationships of casque-headed tree frogs with co-ossified skulls (family Hylidae). University of Kansas Publications. Museum of Natural History, 18, 547–716.

http://biostor.org/reference/51814

Trueb, L. (1974) Systematic relationships of Neotropical horned frogs, genus Hemiphractus (Anura: Hylidae). Occasional Papers of the Museum of Natural History, the University of Kansas, 29, 1–60.

Tyler, M.J. (1994) Australian frogs: a natural history. Reed Books, Chatswood, 192 pp.

Tyler, M.J. & Duellman, W.E. (1995) Superficial mandibular musculature and vocal sac structure in hemiphractine hylid frogs. Journal of Morphology, 224, 65–71.

http://dx.doi.org/10.1002/jmor.1052240108

Valett, B.B. & Jameson, D.L. (1961) The embryology of Eleutherodactylus augusti latrans. Copeia, 1961, 103–109.

http://www.jstor.org/stable/1440182

Varón, A., Vinh, L.S. & Wheeler, W.C. (2010) POY version 4: phylogenetic analysis using dynamic homologies. Cladistics, 26, 72–85.

http://dx.doi.org/10.1111/j.1096-0031.2009.00282.x

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 Linnaean classification schemes. Zootaxa, 3636 (2), 201–244.

http://dx.doi.org/10.11646/zootaxa.3636.2.1

Vera Candioti, M.F. (2005) Morphology and feeding in tadpoles of Ceratophrys cranwelli (Anura: Leptodactylidae). Acta Zoologica, 86, 1–11.

http://dx.doi.org/10.1111/j.0001-7272.2005.00178.x

Warkentin, K.M. (1999) Effects of hatching age on development and hatchling morphology in the red-eyed treefrog, Agalychnis callidryas. Biological Journal of the Linnean Society, 68, 443–470.

http://dx.doi.org/10.1006/bijl.1999.0325

Warkentin, K.M. (2002) Hatching timing, oxygen availability, and external gill regression in the tree frog, Agalychnis callidryas. Physiological and Biochemical Zoology, 75, 155–164.

http://www.jstor.org/stable/10.1086/339214

Wassersug, R.J. & Duellman, W.E. (1984) Oral structures and their development in egg-brooding hylid frog embryos and larvae: evolutionary and ecological significance. Journal of Morphology, 182, l–37.

http://dx.doi.org/10.1002/jmor.1051820102

Weber, L.N. & Caramaschi, U. (2013) A survey of the internal oral morphology in larvae of the genus Hylodes Fitzinger, 1826 (Amphibia, Anura, Hylodidae). Zootaxa, 3635 (5), 557–568.

http://dx.doi.org/10.11646/zootaxa.3635.5.5

Weygoldt, P. (1980) Zur fortpflanzungsbiologie von Phyllobates femora


Refbacks

  • 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