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Type: Article
Published: 2021-12-01
Page range: 351-372
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Description of a new minute frog of the genus Pristimantis (Anura: Strabomantidae) from Cordillera del Condor, Ecuador

Universidad San Francisco de Quito USFQ, Instituto iBIOTROP, Museo de Zoología & Laboratorio de Zoología Terrestre, Quito 170901, Ecuador.
Universidad San Francisco de Quito USFQ, Instituto iBIOTROP, Museo de Zoología & Laboratorio de Zoología Terrestre, Quito 170901, Ecuador.
Universidad San Francisco de Quito USFQ, Instituto iBIOTROP, Museo de Zoología & Laboratorio de Zoología Terrestre, Quito 170901, Ecuador.
Real Jardín Botánico (RJB-CSIC). Plaza de Murillo, 2. 28014 Madrid, Spain.
Museo de Zoología, Escuela de Biología, Pontificia Universidad Católica del Ecuador, Quito, Ecuador
Amphibia Andes Diversity phylogeny Pristimantis daquilemai sp. nov. taxonomy Terrarana


We describe a new species of Pristimantis from southern Ecuador, province of Zamora Chinchipe. The new species is closely related to an undescribed species of Pristimantis from Reserva Tapichalaca, Ecuador and with species of a clade historically assigned to the P. unistrigatus species group, such as P. parvillus, P. luteolateralis, P. walkeri, among others. The new species of Pristimantis is a miniaturized new frog (females 17.1±1.1 mm; males 13.2±0.9 mm), characterized by the presence of “› ‹”-shaped scapular folds, with two subconical tubercles on the medial and posterior regions of folds; tympanic membrane and tympanic annulus present but not externally visible; a prominent rostral papilla present; upper eyelid with one elongated conical tubercle; a conical tubercle on heels; groin with orange or yellow spots. The new species of Pristimantis is distributed in a restricted area in the Cordillera del Condor, a highly-diverse mountain range threatened by multiple anthropogenic activities. We recommend assigning the new species to the Endangered IUCN threatened category because it is only known from three nearby localities within mining concessions.



  1. Almendáriz, A., Brito, J., Batallas, D., Vaca-Guerrero, J. & Ron, S. (2017) Una especie nueva de rana del género Chiasmocleis (Microhylidae: Gastrophryninae) de la Cordillera del Cóndor, Ecuador. Papéis Avulsos de Zoologia, 57, 119–136.
    Almendáriz, A., Brito, J., Batallas, D. & Ron, S. (2014a) Una especie nueva de rana arbórea del género Hyloscirtus (Amphibia: Anura: Hylidae) de la Cordillera del Cóndor. Papéis Avulsos de Zoologia, 54, 33–49.
    Almendáriz, A., Simmons, J.E., Brito, J. & Vaca-Guerrero, J. (2014b) Overview of the herpetofauna of the unexplored Cordillera del Condor of Ecuador. Amphibian & Reptile Conservation, 8, 45–64.
    Almendáriz, A., Ron, S.R. & Brito, J. (2012) Una especie nueva de rana venenosa de altura del género Excidobates (Dendrobatoidea: Dendrobatidae) de la Cordillera del Cóndor. Papéis Avulsos de Zoologia, 52, 387–399.
    Arteaga-Navarro, A.F. & Guayasamin, J.M. (2011) A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the high Andes of Southeastern Ecuador, discovered using morphological and molecular data. Zootaxa, 2876 (1), 17–29.
    Arteaga, A., Pyron, R.A., Penafiel, N., Romero-Barreto, P., Culebras, J., Bustamante, L., Yánez-Muñoz, M. & Guayasamin, M. (2016) Comparative Phylogeography Reveals Cryptic Diversity and Repeated Patterns of Cladogenesis for Amphibians and Reptiles in Northwestern Ecuador. PLoS One, 11 (4), e0151746.
    Arteaga, A., Yanez-Munoz, M. & Guayasamin, J.M. (2013) A new frog of the Pristimantis lacrimosus group (Anura: Craugastoridae) from the montane forests of northwestern Ecuador. Adendum to: The amphibians and reptiles of Mindo: life in the cloudforesty. Serie de Publicaciones Científicas, Universidad Tecnológica Indoamérica, 1, 198–210.
    Barrio-Amoros, C.L., Guayasamín, J.M. & Hedges, B. (2012) A new minute Andean Pristimantis (Anura: Strabomantidae) from Venezuela. Phyllomedusa, 11, 83–93.
    Barrio-Amoros, C.L., Heinicke, M.P. & Hedges, S.B. (2013) A new tuberculated Pristimantis (Anura, Terrarana, Strabomantidae) from the Venezuelan Andes, redescription of Pristimantis pleurostriatus, and variation within Pristimantis vanadisae. Zootaxa, 3647 (1), 43–62.
    Batallas, D. & Brito, J. (2014) Nueva especie de rana del género Pristimantis del grupo lacrimosus (Amphibia: Craugastoridae) del Parque Nacional Sangay, Ecuador. Papéis Avulsos de Zoologia, São Paulo, 54, 51–62.
    Bès de Berc S., Soula, J.C., Baby, P., Souris, M., Christophoul, F. & Rosero, J. (2005) Geomorphic evidence of active deformation and uplift in a modern continental wedge-top–foredeep transition: Example of the eastern Ecuadorian Andes. Tectonophysics, 399, 351–380.
    Boulenger, G.A. (1918) Descriptions of new South American batrachians. Annals and Magazine of Natural History, Series 9, 427–433.
    Brito-Zapata, D. & Reyes-Puig, C. (2021) A new species of terrestrial-breeding frog Pristimantis (Anura: Strabomantidae) from the Cordillera del Cóndor, Zamora Chinchipe, Ecuador. Neotropical Biodiversity, 7 (1), 213–222.
    Brito, J. & Almendáriz, A. (2018) Una especie nueva de rana Pristimantis (Amphibia: Strabomantidae) de ojos rojos de la Cordillera de Cóndor, Ecuador. Cuadernos de Herpetología, 32 (1), 32–40.
    Brito, J., Almendariz, A., Batallas, D. & Ron, S. (2017) Nueva especie de rana bromelícola del género Pristimantis (Amphibia: Craugastoridae), meseta de la cordillera del Cóndor, Ecuador. Papeis Avulsos de Zoologia, Sao Paulo, 57, 177–195.
    Brito, M.J., Ojala-Barbour, R., Batallas, R.D. & Almendáriz, C.A. (2016) A New Species of Pristimantis (Amphibia: Strabomantidae) from the Cloud Forest of Sangay National Park, Ecuador. Journal of Herpetology, 50, 337–344.
    Brito, J., Batallas, R.D. & Velalcázar, D. (2014) Nueva especie de rana terrestre del género Pristimantis (Amphibia: Craugastoridae), meseta de la Cordillera del Cóndor. Papéis Avulsos de Zoologia, 54, 435–466.
    Cairo Carou, H. & Lois, M. (2014) Political Geography of Border Disputes: Changes and Continuities in Latin American Geopolitical Discourse (1990-2013). Cuadernos de Geografía: Revista Colombiana de Geografía, 23, 45–67.
    Canedo, C. & Haddad, C.F. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610–620.
    Cartuche, J.G.T., Romero, C., Carranco, F. & Gramal, A. (2021) Litoestratigrafía, petrografía y geoquímica de la unidad Pachicutza en la cordillera del Cóndor. Revista Científica GeoLatitud, 4 (1), 22–37.
    Chavez, G. & Catenazzi, A. (2016) A new species of frog of the genus Pristimantis from Tingo Maria National Park, Huanuco Department, central Peru (Anura, Craugastoridae). Zookeys, 610, 113–130.
    Chernomor, O., von Haeseler, A. & Minh, B.Q. (2016) Terrace Aware Data Structure for Phylogenomic Inference from Supermatrices. Systematic Biology, 65, 997–1008.
    Cisneros-Heredia, D.F. & Morales-Mite, M. (2008) A new species of glassfrog from elfin forests of the Cordillera del Cóndor, southeastern Ecuador. Herpetozoa, 21 (1/2), 49–56.
    Clark, J.L., Neill, D.A., Weber, A., Gruhn, J.A. & Katan, T. (2010) Shuaria (Gesneriaceae), an Arborescent New Genus from the Cordillera del Cóndor and Amazonian Ecuador. Systematic Botany, 35, 662–674.
    Coltorti, M. & Ollier, C.D. (2000) Geomorphic and tectonic evolution of the Ecuadorian Andes. Geomorphology, 32, 1–19.
    Crawford, A.J., Cruz, C., Griffith, E., Ross, H., Ibanez, R., Lips, K.R., Driskell, A., 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.
    Crawford, A.J., Ryan, M.J. & Jaramillo, C.A. (2010) A new species of Pristimantis (Anura: Strabomantidae) from the Pacific coast of the Darien Province, Panama, with a molecular analysis of its phylogenetic position. Herpetologica, 66 (2), 192–206.
    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.
    De Oliveira, E.A., Rodrigues, L.R., Kaefer, I.L., Pinto, K.C. & Hernandez-Ruz, E.J. (2017) A new species of Pristimantis from eastern Brazilian Amazonia (Anura, Craugastoridae). Zookeys, 687, 101–129.
    De Queiroz, K. (2005) Ernst Mayr and the modern concept of species. Proceedings of the National Academy of Sciences, 102 (suppl 1), 6600–6607.
    De Queiroz, K. (2007) Species concepts and species delimitation. Systematic biology, 56 (6), 879–886.
    Duellman, W.E. & Lehr, E. (2009) Terrestrial-breeding Frogs (Strabomantidae) in Peru. Natur Und Tier-Verlag, Norwich, 384 pp.
    Duellman, W.E. & Wild, E.R. (1993) Anuran amphibians from the Cordillera de Huancabamba, northern Peru: Systematics, ecology, and biogeography. Occasional Papers of the Museum of Natural History, University of Kansas, 157, 1–53.
    Elmer, K.R. & Cannatella, D.C. (2008) Three new species of leaflitter frogs from the upper Amazon forests: cryptic diversity within Pristimantis “ockendeni” (Anura: Strabomantidae) in Ecuador. Zootaxa, 1784 (1), 11–38.
    Elmer, K.R., Davila, J.A. & Lougheed, S.C. (2007) Cryptic diversity and deep divergence in an upper Amazonian leaflitter frog, Eleutherodactylus ockendeni. BMC Evolutionary Biology, 7, 247.
    Espinosa, J., Moreno, J. & Bernal, G. (Eds.) (2018) The Soils of Ecuador. Springer International Publishing, Cham, VII + 164 pp.
    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, 6–228.[0001:SROTFF]2.0.CO;2
    Firkowski, C.R., Bornschein, M.R., Ribeiro, L.F. & Pie, M.R. (2016) Species delimitation, phylogeny and evolutionary demography of co-distributed, montane frogs in the southern Brazilian Atlantic Forest. Molecular Phylogenetics and Evolution, 100, 345–360.
    Foro Intergubernamental sobre Minería, Minerales, Metales y Desarrollo Sostenible (IGF). (2019) Evaluación del Marco de Políticas Mineras del IGF: Ecuador. Winnipeg: IISD.
    Fouquet, A., Noonan, B.P., Rodrigues, M.T., Pech, N., Gilles, A. & Gemmell, N.J. (2012) Multiple quaternary refugia in the eastern Guiana shield revealed by comparative phylogeography of 12 frog species. Systematic Biology, 61 (3), 461–489.
    Frost, D.R. (2021) Amphibian Species of the World: An Online Reference. Version 6.1. American Museum of Natural History, New York, New York. Available from: (accessed 17 November 2021)
    Garcia, J.C., Crawford, A.J., Mendoza, A.M., Ospina, O., Cardenas, H. & Castro, F. (2012) Comparative phylogeography of direct-developing frogs (Anura: Craugastoridae: Pristimantis) in the southern Andes of Colombia. PLoS ONE, 7, E46077.
    Garcia, J.C., Mendoza, A.M., Ospina, O., Cardenas, H. & Castro, F. (2014) A morphometric and molecular approach to define three closely related species of frogs of the Genus Pristimantis (Anura: Craugastoridae) from the Cordillera Occidental in Colombia. Journal of Herpetology, 48, 220–227.
    Gärdenfors, U., Hilton-Taylor, C., Mace, G.M. & Rodríguez, J.P. (2001) The application of IUCN Red List criteria at regional levels. Conservation biology, 15, 1206–1212.
    Glaw, F., Köhler, J., Townsend, T.M. & Vences, M. (2012) Rivaling the world’s smallest reptiles: discovery of miniaturized and microendemic new species of leaf chameleons (Brookesia) from northern Madagascar. PLoS ONE, 7 (2), e31314.
    Gorin, V.A., Scherz, M.D., Korost, D.V. & Poyarkov, N.A. (2021) Consequences of parallel miniaturisation in Microhylinae (Anura, Microhylidae), with the description of a new genus of diminutive South East Asian frogs. Zoosystematics and Evolution, 97 (21), 21–54.
    Guayasamin, J.M., Hutter, C.R., Tapia, E.E., Culebras, J., Penafiel, N., Pyron, R.A., Morochz, C., Funk, C. & Arteaga, A. (2017) Diversification of the rainfrog Pristimantis ornatissimus in the lowlands and Andean foothills of Ecuador. PLoS ONE, 12, e0172615.
    Guayasamin, J.M., Krynak, T., Krynak, K., Culebras, J. & Hutter, C.R. (2015) Phenotypic plasticity raises questions for taxonomically important traits: a remarkable new Andean rainfrog (Pristimantis) with the ability to change skin texture. Zoological Journal of the Linnean Society, 173, 913–928.
    Guindon, S., Dufayard, J.F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321.
    Hanken, J. & Wake, D.B. (1993) Miniaturization of body size: organismal consequences and evolutionary significance. Annual Review of Ecology and Systematics, 24, 501–519.
    Hedges, S.B. (1994) Molecular Evidence For The Origin Of Birds. Proceedings of the National Academy of Sciences of the United States of America, 91, 2621–2624.
    Hedges, S.B., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737 (1), 1–181.
    Heinicke, M.P., Barrio-Amoros, C.L. & Hedges, S.B. (2015) Molecular and morphological data support recognition of a new genus of New World direct-developing frog (Anura: Terrarana) from an under-sampled region of South America. Zootaxa, 3986 (2), 151–172.
    Heinicke, M.P., Duellman, W.E., Trueb, L., Means, D.B., MacCulloch, R.D. & Hedges, B. (2009) A new frog family (Anura: Terrarana) from South America and an expanded direct-eeveloping clade revealed by molecular phylogeny. Zootaxa, 2211 (1), 1–35
    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 of the United States of America, 104 (24), 10092–10097.
    Heinicke, M.P., Lemmon, A.R., Lemmon, E.M., McGrath, K. & Hedges, S.B. (2018) Phylogenomic support for evolutionary relationships of New World direct-developing frogs (Anura: Terraranae). Molecular Phylogenetics and Evolution, 118, 145–155.
    Ibarra, H.G.R. (2018) La rebelión de Daquilema (Yaruquíes-Chimborazo, 1871). Colección Testigos de la Historia. Cooperativa Fernando Daquilema, Instituto Nacional de Patrimonio Cultural, Quito, 84 pp.
    IGM (2012) Mapa Geográfico República del Ecuador. Instituto Geográfico Militar, Quito. Availabel from: (accessed 16 November 2021)
    Jablonski, D., Grula, D., Barrio-Amoros, C.L. & Kok, P.J.R. (2017) Molecular phylogenetic relationships among Pristimantis summit populations in the eastern tepui chain: insights from P. aureoventris (Anura: Craugastoridae). Salamandra, 53, 473–478.
    Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589.
    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.
    Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.
    Kok, P.J., Dezfoulian, R., Means, D.B., Fouquet, A. & Barrio-Amoros, C. (2018) Amended diagnosis and redescription of Pristimantis marmoratus (Boulenger, 1900) (Amphibia: Craugastoridae), with a description of its advertisement call and notes on its breeding ecology and phylogenetic relationships. European Journal of Taxonomy, 397, 1–30.
    Kok, P.J., MacCulloch, R.D., Means, D.B., Roelants, K., Van Bocxlaer, I. & Bossuyt, F. (2012) Low genetic diversity in tepui summit vertebrates. Current Biology, 22, R589–R590.
    Lehr, E., Moravec, J., Cusi, J.C. & Gvozdik, V. (2017) A new minute species of Pristimantis (Amphibia: Anura: Craugastoridae) with a large head from the Yanachaga-Chemillen National Park in central Peru, with comments on the phylogenetic diversity of Pristimantis occurring in the Cordillera Yanachaga. European Journal of Taxonomy, 325, 1–22.
    Lehr, E., von May, R., Moravec, J. & Cusi, J.C. (2017) Three new species of Pristimantis (Amphibia, Anura, Craugastoridae) from upper montane forests and high Andean grasslands of the Pui Pui Protected Forest in central Peru. Zootaxa, 4299 (3), 301–336.
    Lehr, E., Moravec, J. & Cusi, J.C. (2012) Two new species of Phrynopus (Anura, Strabomantidae) from high elevations in the Yanachaga-Chemillen National park in Peru (Departamento de Pasco). Zookeys, 235, 51–71.
    Lehr, E. & Coloma, L.A. (2008) A minute new ecuadorian Andean frog (Anura: Strabomantidae, Pristimantis). Herpetologica, 64, 354–367.
    Lehr, E., Aguilar, C., Siu-Ting, K. & Carlos Jordán, J. (2007) Three New Species of Pristimantis (Anura: Leptodactylidae) from the Cordillera De Huancabamba in Northern Peru. Herpetologica, 63 (4), 519–536.[519:TNSOPA]2.0.CO;2
    Lessmann, J., Troya, M.J., Flecker, A.S., Funk, W.C., Guayasamin, J.M., Ochoa-Herrera, V., Poff, N., Suárez, E. & Encalada, A. (2019) Validating anthropogenic threat maps as a tool for assessing river ecological integrity in Andean-Amazon basins. PeerJ, 7, e8060.
    López-Ocón, L. (1986) Etnogénesis y rebeldía andina: la sublevación de Fernando Daquilema en la provincia del Chimborazo en 1871. Boletín americanista, 36, 113–133.
    Lynch, J.D. & Duellman, W.E. (1997) Frogs of the Genus Eleutherodactylus in Western Ecuador. University of Kansas, Natural History Museum, Special Publication 23. Natural History Museum, University of Kansas, Lawrence, Kansas, 236 pp.
    Lynch, J.D. & Duellman, W.E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). The University of Kansas, Museum of Natural History, Miscellaneous Publications, 69, 1–86.
    Lynch, J.D. (1974) A new species of Eleutherodactylus (Amphibia: Leptodactylidae) from the Pacific lowlands of Ecuador. Proceedings of the Biological Society of Washington, 87, 381–388.
    Lynch, J.D. (1976) New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador. Occasional Papers of the Museum of Natural History, University of Kansas, 55, 1–33.
    Maddison, W.P. & Maddison, D.R. (2019) Mesquite: a modular system for evolutionary analysis. Version 3.61. [program]
    Monteiro, J.P.C., Condez, T.H., García, P.C.A., Comitti, E.J., Amaral, I.B. & Haddad, C.F. (2018) A new species of Brachycephalus (Anura, Brachycephalidae) from the coast of Santa Catarina State, southern Atlantic Forest, Brazil. Zootaxa, 4407 (4), 483–505.
    Morales, C., Jadán, O. & Aguirre, Z. (2013) Bosque siempreverde montano de las cordilleras del Cóndor-Kutukú. In: Sistema de Clasificación de Ecosistemas del Ecuador Continental, Quito, 2013, pp. 205–206.
    Mutlak, N.D. (2009) Daquilema, Fernando (d. 1872) and the 1871 uprising, Ecuador. Available from: (accessed 16 November 2021)
    Neill, D. (2005) Cordillera del Cóndor: botanical treasures between the Andes and the Amazon. PlantTalk, 41, 17–21.
    Neill, D.A. (2007) MBG: Research: Ecuador: Botanical Exploration of the Cordillera del Cóndor Region. Botanical Exploration of the Cordillera del Cóndo. Available from: (accessed 31 March 2019)
    Nguyen, L.T., Schmidt, H.A., von Haeseler, A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution, 32, 268–275.
    Ortega-Andrade, H.M. & Venegas, P.J. (2014) A new synonym for Pristimantis luscombei (Duellman and Mendelson 1995) and the description of a new species of Pristimantis from the upper Amazon basin (Amphibia: Craugastoridae). Zootaxa, 3895 (1), 31–57.
    Pabijan, M., Wollenberg, K.C. & Vences, M. (2012) Small body size increases the regional differentiation of populations of tropical mantellid frogs (Anura: Mantellidae). Journal of Evolutionary Biology, 25, 2310–2324.
    Padial, J.M., Chaparro, J.C., Castroviejo-Fisher, S., Guayasamín, J.M., Lehr, E., Delgado, A.J., Vaira, M., Teixeira, M., Aguayo, 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.
    Paez, N.B. & Ron, S.R. (2019) Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species. Zookeys, 868, 1–112.
    Paz, A., Ibanez, R., Lips, K.R. & Crawford, A.J. (2015) Testing the role of ecology and life history in structuring genetic variation across a landscape: a trait-based phylogeographic approach. Molecular Ecology, 24, 3723–3737.
    Pinto-Sanchez, N.R., Ibanez, R., Madrinan, S., Sanjur, O.I., Bermingham, E. & Crawford, A.J. (2012) The Great American Biotic Interchange in frogs: multiple and early colonization of Central America by the South American genus Pristimantis (Anura: Craugastoridae). Molecular Phylogenetics and Evolution, 62, 954–972.
    Ramírez Jaramillo, S., Reyes-Puig, C.P., Batallas, D. & Yánez-Muñoz, M.H. (2018) Ranas terrestres en los ecosistemas surandinos de Ecuador IV: Una nueva especie de Pristimantis (Anura: Strabomantidae) de los páramos sur del Parque Nacional Sangay. Avances en Ciencias e Ingenierías, 10, 1–17.
    Rivera-Correa, M. & Daza, J.M. (2016) Molecular phylogenetics of the Pristimantis lacrimosus species group (Anura: Craugastoridae) with the description of a new species from Colombia. Acta Herpetologica, 11, 31–45.
    Rivera-Prieto, D.A., Rivera-Correa, M. & Daza, J.M. (2014) A new colorful species of Pristimantis (Anura: Craugastoridae) from the eastern flank of the Cordillera Central in Colombia. Zootaxa, 3900 (2), 223–242.
    Roddaz,. M., Christophoul, F., Burgos Zambrano, J.D., Soula, J.C. & Baby, P. (2012) Provenance of late Oligocene to quaternary sediments of the Ecuadorian Amazonian foreland basin as inferred from major and trace element geochemistry and Nd–Sr isotopic composition. Journal of South American Earth Sciences, 37, 136–153.
    Ron, S.R., Merino-Viteri, A. & Ortiz, D.A. (2021) Anfibios del Ecuador. Version 2021.0. Museo de Zoología, Pontificia Universidad Católica del Ecuador, Quito. [program]
    Ron, S.R., Caminer, M.A., Varela-Jaramillo, A. & Almeida-Reinoso, D. (2018) A new treefrog from Cordillera del Cóndor with comments on the biogeographic affinity between Cordillera del Cóndor and the Guianan Tepuis (Anura, Hylidae, Hyloscirtus). ZooKeys, 809, 97–124.
    Roy, B.A., Zorrilla, M., Endara, L., Thomas, D.C., Vandegrift, R., Rubenstein, J.M., Policha, T., Ríos-Touma, B. & Read, M. (2018) New mining concessions could severely decrease biodiversity and ecosystem services in Ecuador. Tropical Conservation Science, 11, 1–20.
    Shepack, A., von May, R., Ttito, A. & Catenazzi, A. (2016) A new species of Pristimantis (Amphibia, Anura, Craugastoridae) from the foothills of the Andes in Manu National Park, southeastern Peru. ZooKeys, 594, 143–164.
    Schulenberg, T.S. & Awbrey, K. (Eds.) (1997) The Cordillera del Condor Region of Ecuador and Peru. Conservation International, Washington, D.C., 231 pp. Available from: (accessed 1 April 2019)
    Szekely, P., Cogalniceanu, D., Szekely, D., Paez, N. & Ron, S.R. (2016) A new species of Pristimantis from southern Ecuador (Anura, Craugastoridae). Zookeys, 606, 77–97.
    Terán, F. (1984) Geografía del Ecuador. 11th Edition. Libresa, Quito, 462 pp.
    Terán-Valdez, A. & Guayasamin, J.M. (2010) The smallest terrestrial vertebrate of Ecuador: A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the Cordillera del Cóndor. Zootaxa, 2447 (1), 53–68.
    Urgilés, V.L., Székely, P., Székely, D., Christodoulides, N., Sanchez-Nivicela, J.C. & Savage, A.E. (2019) Genetic delimitation of Pristimantis orestes (Lynch, 1979) and P. saturninoi Brito et al., 2017 and description of two new terrestrial frogs from the Pristimantis orestes species group (Anura, Strabomantidae). Zookeys, 864, 111–146.
    von May, R., Catenazzi, A., Corl, A., Santa-Cruz, R., Carnaval, A.C. & Moritz, C. (2017) Divergence of thermal physiological traits in terrestrial breeding frogs along a tropical elevational gradient. Ecology and Evolution, 7 (9), 3257–3267.
    Waddell, E.H., Crotti, M., Lougheed, S.C., Cannatella, D.C. & Elmer, K.R. (2018) Hierarchies of evolutionary radiation in the world’s most species rich vertebrate group, the Neotropical Pristimantis leaf litter frogs. Systematics and Biodiversity, 16, 807–819.
    Wassersug, R.J., Yang, H., Sepkoski, J.J. & Raup, D.M. (1979) The evolution of body size on islands: a computer simulation. The American Naturalist, 114, 287–295.
    Wiens, J.J. & Graham, C.H. (2005) Niche conservatism: integrating evolution, ecology, and conservation biology. Annual Review of Ecology Evolution and Systematics, 36, 519–539.
    Wiens, J.J., Ackerly, D.D., Allen, A.P., Anacker, B.L., Buckley, L.B., Cornell, H.V., Damschen, E.I., Jonathan Davies, T., Grytnes, J.A., Harrison, S.P., Hawkins, B.A., Holt, R.D., McCain, C.M. & Stephens, P.R. (2010) Niche conservatism as an emerging principle in ecology and conservation biology. Ecology Letters, 13, 1310–1324.
    Winckell, A., Zebrowski, C. & Sourdat, M. (Eds.) (1997) Los paisajes naturales del Ecuador: Las regiones y paisajes del Ecuador. Centro Ecuatoriano de Investigación Geográfica, Instituto Panamericano de Geografía e Historia, Instituto Geografico Militar, Instituto Francés de Investigación Científica para el Desarrollo en Cooperación ORSTOM, Quito, 417 pp.
    Wollenberg, K.C., Vieites, D.R., Van der Meijden, A., Glaw, F., Cannatella, D.C. & Vences, M. (2008) Patterns of endemism and species richness in Malagasy cophyline frogs support a key role of mountainous areas for speciation. Evolution, 62, 1890–1907.
    Wollenberg, K.C., Vieites, D.R., Glaw, F. & Vences, M. (2011) Speciation in little: the role of range and body size in the diversification of Malagasy mantellid frogs. BMC Evolutionary Biology, 11, 217.
    Womack, M.C. & Bell, R.C. (2020) Two-hundred million years of anuran body-size evolution in relation to geography, ecology and life history. Journal of Evolutionary Biology, 33, 1417–1432.
    Yeh, J. (2002) The effect of miniaturized body size on skeletal morphology in frogs. Evolution, 56, 628–641.
    Zhang, P., Liang, D., Mao, R.L., Hillis, D.M., Wake, D.B. & Cannatella, D.C. (2013) Efficient sequencing of Anuran mtDNAs and a mitogenomic exploration of the phylogeny and evolution of frogs. Molecular Biology and Evolution, 30, 1899–1915.