Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2022-02-22
Page range: 521-540
Abstract views: 948
PDF downloaded: 41

A new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Strabomantidae) from the “Brejos de Altitude” in Northeast Brazil

Departamento de Biologia, Centro de Ciências, Campus do Pici, Universidade Federal do Ceará, Fortaleza, Ceará, Brazil.
Laboratório de Vertebrados, Instituto de Ciências Biológicas, Universidade Federal do Rio Grande, Av. Itália, Km 8, 96203-900, Rio Grande, Rio Grande do Sul, Brazil
Departamento de Biodiversidade e Centro de Aquicultura (CAUNESP), Instituto de Biociências, Universidade Estadual Paulista (UNESP), Av. 24A, 1515, Rio Claro, São Paulo, CEP 13506-900, Brazil
Departamento de Biodiversidade e Centro de Aquicultura (CAUNESP), Instituto de Biociências, Universidade Estadual Paulista (UNESP), Av. 24A, 1515, Rio Claro, São Paulo, CEP 13506-900, Brazil
Departamento de Biologia, Centro de Ciências, Campus do Pici, Universidade Federal do Ceará, Fortaleza, Ceará, Brazil.
Amphibia Ceará State Neotropics Systematics Taxonomy

Abstract

Pristimantis is the most diverse Neotropical genus of terrestrial vertebrates and is distributed from Central America to Argentina. The last few years have seen the description of several new species of the genus, suggesting that its diversity is still underestimated. After decades of uncertainties about the taxonomic status of populations of Pristimantis from the “Brejos de Altitude” of the state of Ceará, Northeast Brazil, we finally found morphological, acoustic and molecular evidence confirming their distinctiveness from other Atlantic Forest species. The new species is characterized by the following: shagreen dorsal skin with small-scattered tubercles, absence of dorsolateral fold, presence of tarsal fold, advertisement call composed of 1–8 pulsed notes, (2–5 pulses per note), and dominant frequency located in the second energy band, ranging from 3617–4220 Hz. Phylogenetic analysis placed the new species in the Pristimantis conspicillatus species group and the sister lineage of the Atlantic Forest clade comprising P. ramagii, P. paulodutrai and the P. vinhai species complex.

 

References

  1. Acevedo-Rincón, A.A., Armesto, O. & Palma, R.E. (2020) Two new species of Pristimantis (Anura: Craugastoridae) with notes on the distribution of the genus in northeastern Colombia. Zootaxa, 4750 (4), 499–523. https://doi.org/10.11646/zootaxa.4750.4.3
    Barbour, T. & Noble, G.K. (1920) Some amphibians from northwestern Peru, with a revision of the genera Phyllobates and Telmatobius. Bulletin of the Museum of Comparative Zoology at Harvard, 63, 395–427.
    Brown, S.D.J., Collins, R.A., Boyer, S., Lefort, M.C., Malumbresolarte, J., Vink, C.J. & Cruickshank, R.H. (2012) Spider: a R package for the analysis of species identity and evolution, with particular reference to DNA barcoding. Molecular Ecology Resources, 12, 562–565. https://doi.org/10.1111/j.1755-0998.2011.03108.x
    Bokermann, W.C.A. (1975) Três espécies novas de Eleutherodactylus do sudeste da Bahia, Brasil (Anura, Leptodactylidae). Revista Brasileira de Biologia, 34 (1), 11–18.
    Borges-Nojosa, D.M. & Caramaschi, U. (2003) Composição e análise comparativa da diversidade e das afinidades biogeográficas dos lagartos e anfisbenídeos (Squamata) dos brejos nordestinos. In: Leal, I.R., Tabarelli, M. & Silva, J.M.C. (Eds.), Ecologia e Conservação da Caatinga. Ed. Universitária, Universidade Federal de Pernambuco, Recife, pp. 463–512.
    Borges-Nojosa, D.M., Caramaschi, U. & Rodrigues, M.T. (2016) A new species of lizard Placosoma Tschudi, 1847 (Squamata: Gymnophthalmidae) from the relictual forest mountains of the State of Ceará, Brazil. Zootaxa, 4169 (1), 160–170. https://doi.org/10.11646/zootaxa.4169.1.8
    Borges-Nojosa, D.M. & Lima, D.C. (2001) Dieta de Drymoluber dichrous (Peters, 1863) dos Brejos-de-Altitude do estado do Ceará, Brasil (Serpentes, Colubridae). Boletim do Museu Nacional, Nova Serie, Zoologia, 468, 1–5.
    Boulenger, G.A. (1888) On some Reptiles and Batrachians from Iguarasse, Pernambuco. The Annals and Magazine of Natural History, Series 6, 2 (7), 40–43. https://doi.org/10.1080/00222938809460873
    Boulenger, G.A. (1898) An account of the reptiles and batrachians collected by Mr. F. H. Rosenberg in western Ecuador. Proceedings of the Zoological Society of London, 1898, 107–126.
    Canedo, C. & Haddad, C.F.B. (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 (2), 610–620. https://doi.org/10.1016/j.ympev.2012.07.016
    Carnaval, A.C. & Bates, J.M. (2007) Amphibian DNA shows marked genetic structure and tracks Pleistocene climate change in Northeastern Brazil. Evolution, 61 (12), 2942–2957. https://doi.org/10.1111/j.1558-5646.2007.00241.x
    Castro, D.P., Mângia, S., Medeiros-Magalhães, F., Röhr, D.L., Camurugi, F., Silveira-Filho, R.R., Silva, M.M.X., Andrade-Oliveira, J.A., Sousa, T.A., França, F.G.R., Harris, D.J., Garda, A.A. & Borges-Nojosa, D.M. (2019) Herpetofauna of protected areas in the Caatinga VI: The Ubajara National Park, Ceará, Brazil. Herpetology Notes, 12, 727–742.
    Center for Conservation Bioacustics. (2019) Raven Pro: interactive sound analysis software, Version 1.6.1. The Cornell Lab of Ornithology, Ithaca, New York. Available from: http//www.birds.cornell.edu/raven (accessed 10 January 2021)
    Cocroft, R.B & Ryan, M.J. (1995) Patterns of advertisement call evolution in toads and chorus frogs. Animal Behaviour, 49 (2), 283–303. https://doi.org/10.1006/anbe.1995.0043
    Duellman, W.E. (1992) A new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from northeastern Peru. Revista Española de Herpetología, 6, 23–29.
    Duellman, W.E. & Hedges, S.B. (2005) Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera Yanachaga in Central Peru. Copeia, 2005 (3), 526–538. https://doi.org/10.1643/CH-05-019R
    Duellman, W.E. & Hedges, S.B. (2007) Three new species of Pristimantis (Lissamphibia, Anura) from montane forests of the Cordillera Yanachaga in Central Peru. Phyllomedusa, 6 (2), 119–135. https://doi.org/10.11606/issn.2316-9079.v6i2p119-135
    Duellman, W.E. & Lehr, E. (2007) Frogs of the genus Eleutherodactylus (Leptodactylidae) in the Cordillera Occidental in Peru with descriptions of three new species. Scientific Papers, Natural History Museum, The University of Kansas, 39, 1–13.
    Duellman, W.E. & Pramuk, J.B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers, Natural History Museum, The University of Kansas, 13, 1–78.
    Dunn, E.R. (1931) New frogs from Panama and Costa Rica. Occasional Papers of the Boston Society of Natural History, 5, 385–401.
    Feller, A.E. & Hedges, S.B. (1998) Molecular evidence for the early history of living amphibians. Molecular Phylogenetics and Evolution, 9 (3), 509–516. https://doi.org/10.1006/mpev.1998.0500
    Flores, G. & Rodríguez, L.O. (1997) Two new species of the Eleutherodactylus conspicillatus group (Anura: Leptodactylidae) from Peru. Copeia, 1997 (2), 388–394. https://doi.org/10.2307/1447759
    Fouquet, A., Loebmann, D., Castroviejo-Fisher, S., Padial, J.M., Orrico, V.G.D., Lyra, M.L., Roberto, I.J., Kok, P.J.R., Haddad, C.F.B. & Rodrigues, M.T. (2012) From Amazonia to the Atlantic forest, Molecular phylogeny of Phyzelaphryninae frogs reveals unexpected diversity and a striking biogeographic pattern emphasizing conservation challenges. Molecular Phylogenetics and Evolution, 65 (2), 547–561. https://doi.org/10.1016/j.ympev.2012.07.012
    Frost, D.R. (2021) Amphibian Species of the World, an Online Reference. Version 6.1. American Museum of Natural History, New York. Available from http://research.amnh.org/herpetology/amphibia/index.html (accessed 2 February 2022)
    Giaretta, A.A., Teixeira, B.F.V., Bernardes, C.S. & Marinho, P. (2018) Distribution, call and habitat of Pristimantis dundeei (Anura, Craugastoridae). Neotropical Biodiversity, 4 (1), 134–137. https:// doi.org/https://doi.org/10.1080/23766808.2018.1506274
    González-Durán, G.A., Targino, M., Rada, M. & Grant, T. (2017) Phylogenetic relationships and morphology of the Pristimantis leptolophus species group (Amphibia: Anura: Brachycephaloidea), with the recognition of a new species group in Pristimantis Jiménez de la Espada, 1870. Zootaxa, 4243 (1), 42–74. https://doi.org/10.11646/zootaxa.4243.1.2
    Günther, A. (1858) Neue Batrachier in der Sammlung des Britischen Museums. Archiv für Naturgeschichte, 24, 319–328.
    Hedges, S.B. (1994) Molecular evidence for the origin of birds. Proceedings of the National Academy of Science, 91 (7), 2621–2624. https://doi.org/10.1073/pnas.91.7.2621
    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), 1–182. https://doi.org/10.11646/zootaxa.1737.1.1
    Heyer, W.R. & Barrio-Amorós, C.L. (2009) The advertisement calls of two sympatric frogs, Leptodactylus lithonaetes (Amphibia: Anura: Leptodactylidae) and Pristimantis vilarsi (Amphibia: Anura: Strabomantidae). Proceedings of the Biological Society of Washington, 122 (3), 282–291. https://doi.org/10.2988/09-02.1
    Heyer, W.R. & De Carvalho, C.M. (2000) The enigmatic advertisement call of Eleutherodactylus ramagii (Amphibia: Anura: Leptodactylidae). Amphibia-Reptilia, 21, 117–121.
    Heyer, W.R. & Muñoz, A.M. (1999) Validation of Eleutherodactylus crepitans Bokermann, 1965, notes on the types and type locality of Telatrema heterodactylum Miranda-Ribeiro, 1937, and a description of a new species of Eleutherodactylus from Mato Grosso, Brazil. Proceedings of the Biological Society of Washington, 112, 1–18.
    Heyer, W.R., Rand, A.S., Cruz, C.A.G., Peixoto, O.L. & Nelson, C.E. (1990) Frogs of Boracéia. Arquivos de Zoologia, 31 (4), 231–410. https://doi.org/10.11606/issn.2176-7793.v31i4p231-410
    Hoogmoed, M.S., Borges, D.M. & Cascon, P. (1994) Three new species of the genus Adelophryne (Amphibia: Anura: Leptodactylidae) from northeastern Brazil, with remarks on the other species of the genus. Zoologische Mededelingen, 68, 271–300.
    Hoogmoed, M.S., Lynch, J.D. & Lescure, J. (1977) A new species of Eleutherodactylus from Guiana (Leptodactylidae, Anura). Zoologische Mededelingen, 51, 33–42.
    Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software Version 7: improvements in performance and usability. Molecular Biology and Evolution, 30 (4), 772–780. https:// doi.org/10.1093/molbev/mst010
    Kaiser, H., Dwyer, C.M., Feichtinger, W. & Schmid, M. (1995) A new species of Eleutherodactylus (Anura: Leptodactylidae) from Tobago, West Indies, and its morphometric and cytogenetic characterization. Herpetological Natural History, 3 (2), 151–163.
    Köhler, J., Jansen, M., Rodríguez, A., Kok, P.J.R., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F.B., Rodel, M.O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251 (1), 1–124. https://doi.org/10.11646/zootaxa.4251:1–124
    Köhler, J. & Jungfer, K.H. (1995) Eine neue Art und ein Erstnachweis von Fröschen der Gattung Eleutherodactylus aus Bolivien. Salamandra, 31 (3), 149–156.
    Lehr, E. & Duellman, W.E. (2007) Two new species of Eleutherodactylus (Anura: Leptodactylidae) from the Cordillera Occidental in Peru. Copeia, 2007 (1), 140–149. https://doi.org/10.1643/0045-8511(2007)7[140:TNSOEA]2.0.CO;2
    Lima-Verde, J.S. & Cascon, P. (1990) Lista preliminar da herptofauna do Ceará, Brasil. Caatinga, 7, 158–163.
    Loebmann, D. & Haddad, C.F.B. (2010) Amphibians and reptiles from a highly diverse area of the Caatinga domain: composition and conservation implications. Biota Neotropica, 10 (3), 228–256. https://doi.org/10.1590/S1676-06032010000300026
    Lynch, J.D. (1980) A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates, 2696, 1–24.
    Lynch, J.D. (1994) Two new species of the Eleutherodactylus conspicillatus group (Amphibia: Leptodactylidae) from the Cordillera Oriental of Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 19 (72), 187–193.
    Lynch, J.D. & Duellman, W.E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 69, 1–86. https://doi.org/10.5962/bhl.title.16222
    Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador: Systematics, Ecology, and Biogeography. Natural History Museum University Kansas, Lawrence, Kensas, 236 pp. https://doi.org/10.5962/bhl.title.7951
    Lynch, J.D. & Hoogmoed, M.S. (1977) Two new species of Eleutherodactylus (Amphibia: Leptodactylidae) from northeastern South America. Proceedings of the Biological Society of Washington, 90, 424–439.
    Lyra, M.L., Haddad, C.F.B. & de Azeredo-Espin, A.M.L. (2017) Meeting the challenge of DNA barcoding Neotropical amphibians: polymerase chain reaction optimization and new COI primers. Molecular Ecology Resources, 17 (5), 966–980. https://doi.org/10.1111/1755- 0998.12648
    Lutz, A. (1927) Notas sobre batrachios da Venezuela e da Ilha de Trinidad/Notes on batrachians from Venezuela and Trinidad. Memórias do Instituto Oswaldo Cruz, Rio de Janeiro, 20, 35–65.
    Maciel, N.M., Vaz-Silva, W., de Oliveira, R.M. & Padial, J.M. (2012) A new species of Pristimantis (Anura: Strabomantidae) from the Brazilian Cerrado. Zootaxa, 3265 (1), 43–56. https://doi.org/10.11646/zootaxa.3265.1.3
    Malagoli, L.R., Pezzuti, T.L., Bang, D.L., Faivovich, J., Lyra, M.L., Giovanelli, J.G.R., Garcia, P.C.A., Sawaya, R.J. & Haddad, C.F.B. (2021) A new reproductive mode in anurans: Natural history of Bokermannohyla astartea (Anura: Hylidae) with the description of its tadpole and vocal repertoire. PLoS One, 16, e0246401. https://doi.org/10.1371/journal.pone.0246401
    Mângia, S., Koroiva, R., Nunes, P.M.S., Roberto, I.J., Ávila, R.W., Sant’Anna, A.C., Santana, D.J. & Garda, A.A. (2018) A new species of Proceratophrys (Amphibia: Anura: Odontophrynidae) from the Araripe Plateau, Ceará State, northeastern Brazil. Herpetologica, 74 (3), 255–268. https://doi.org/10.1655/Herpetologica-D-16-00084.1
    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. 2010 Gateway Computing Environments Workshop (GCE), 2010, 1–8. https://doi.org/10.1109/GCE.2010.5676129
    Melin, D.E. (1941) Contributions to the knowledge of the Amphibia of South America. Göteborgs Kungl. Vetenskaps-och Vitterhets-samhälles. Handlingar. Serien B, Matematiska och Naturvetenskapliga Skrifter, 1, 1–71.
    Myers, C.W. & Duellman, W.E. (1982) A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Novitates, 2752, 1–32.
    Oitaven, L.P.C., Santos, J.R.O., Silva, A.D.O., Gambale, P.G. & Moura, J.G.B. (2017) Description of vocalisations and analysis of intra and inter-individual variation in Pristimantis ramagii (Boulenger, 1888) in an upland swamp, northeast Brazil. Herpetology Notes, 10, 197–203.
    Oliveira, E.A., Rodrigues, L.R., Kaefer, I.L., Pinto, K.C., Hernández-Ruiz, E.J. (2017) A new species of Pristimantis from eastern Brazilian Amazonia (Anura, Craugastoridae). ZooKeys, 687, 101–129. https://doi.org/10.3897/zookeys.687.13221
    Oliveira, E.A., Silva, L.A., Guimarães, K.L.A., Penhacek, M., Martínez, J.G., Rodrigues, L.R.R., Santana, D.J. & Hernández-Ruz, E.J. (2020) Four new species of Pristimantis Jiménez de la Espada, 1870 (Anura: Craugastoridae) in the eastern Amazon. PLoS One, 15, 1–28. https://doi.org/10.1371/journal.pone.0229971
    Padial, J.M. & de la Riva, I. (2009) Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura). Zoological Journal of the Linnean Society, 155 (1), 97–122. https://doi.org/10.1111/j.1096-3642.2008.00424.x
    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. https://doi.org/10.11646/zootaxa.3825.1.1
    Padial, J.M., Gagliardi-Urrutia, G., Chaparro, J.C. & Gutiérrez, R.C. (2016) A new species of the Pristimantis conspicillatus Group from the Peruvian Amazon (Anura: Craugastoridae). Annals of Carnegie Museum, 83 (3), 207–218. https://doi.org/10.2992/007.083.0302
    Palumbi, S.R., Martin, A.P., Kessing, B.D. & Mcmillan, W.O. (1991) Detecting population structure using mitochondrial DNA. In: Hoelzel, A.R. (Ed.), Genetic Ecology of Whales and Dolphins. International Whaling Commission Special Issue 13. International Whaling Commission, Cambridge, pp. 203–215.
    R Development Core Team. (2019) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available from: https://www.r-project.org/ (accessed 16 July 2021)
    Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
    Rivero, J.A. & Serna, M.A. (1988) Tres nuevas especies de Eleutherodactylus (Amphibia, Leptodactylidae) de Antioquia, Colombia. Caribbean Journal of Science, 23, 386–399.
    Roberto, I.J. & Loebmann, D. (2016) Composition, distribution patterns, and conservation priority areas for the herpetofauna of the state of Ceará, northeastern Brazil. Salamandra, 52, 134–152.
    Roberto, I.J., Oliveira, C.R.D., Araújo-Filho, J.A.D., Oliveira, H.F.D. & Ávila, R.W. (2017) The herpetofauna of the Serra do Urubu mountain range: a key biodiversity area for conservation in the Brazilian Atlantic forest. Papéis Avulsos de Zoologia, 57 (27), 347–373. https://doi.org/10.11606/0031-1049.2017.57.27
    Rocha, D. (1948) Subsídio para o estudo da fauna cearense (Catálogo das espécies de animais por mim coligidas e anotadas). Revista do Instituto do Ceará, 62, 102–138.
    Rodriguez, L. (1994) A new species of the Eleutherodactylus conspicillatus Group from Peru, with comments on its call. Alytes, 12, 49–63.
    Ronquist, F., Teslenko, M., Van der mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mrbayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. https://doi.org/10.1093/sysbio/sys029
    Santos, A.M.M., Cavalcanti, D.R., da Silva, J.M.C. & Tabarelli, M. (2007) Biogeographical relationships among tropical forests in north-eastern Brazil. Journal of Biogeography, 34, 437–446. https://doi.org/10.1111/j.1365-2699.2006.01604.x
    Savage, J.M. & Heyer, W.R. (1967) Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America. Beiträge zur Neotropischen Fauna, 5, 111–131. https://doi.org/10.1080/01650526709360400
    Silva, J.M.C. & Castelleti, C.H.M. (2003) Status of the biodiversity of the Atlantic Forest of Brazil. In: Galindo-Leal, C. & Camara, I.G. (Eds.), The Atlantic Forest of South America: Biodiversity Status, Threats, and Outlook. CABS and Island Press, Washington, pp. 31–42.
    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313. https://doi.org/10.1093/bioinformatics/btu033
    Steindachner, F. (1864) Batrachologische Mittheilungen. Verhandlungen des Zoologisch-Botanischen Vereins in Wien, 14, 239–288.
    Sueur, J., Aubin, T. & Simonis, C. (2008) Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18, 213–226. https://doi.org/10.1080/09524622.2008.9753600
    Taucce, P.P.G., Nascimento, J.S., Trevisan, C.C., Leite, F.S.F., Santana, D.J., Haddad, C.F.B. & Napoli, M.F. (2020) A new rupicolous species of the Pristimantis conspicillatus group (Anura: Brachycephaloidea: Craugastoridae) from central Bahia, Brazil. Journal of Herpetology, 54 (2), 245–257. https://doi.org/10.1670/19-114
    Trevisan, C.C., Batalha-Filho, H., Garda, A.A., Menezes, L., Dias, I.R., Sol, M., Canedo, C., Junc, F.A. & Napoli, M.F. (2020) Cryptic diversity and ancient diversification in the northern Atlantic Forest Pristimantis (Amphibia, Anura, Craugastoridae). Molecular Phylogenetics and Evolution, 148, 1055–7903. https://doi.org/10.1016/j.ympev.2020.106811
    Ttito, A. & Catenazzi, A. (2021) Pristimantis achupalla sp. n., a new minute species of direct-developing frog (Amphibia, Anura, Strabomantidae) inhabiting bromeliads of the montane forest of the Amazonian Andes of Puno, Peru. PeerJ, 9, e11878. https://doi.org/10.7717/peerj.11878
    Zumel, D., Buckley, D. & Ron, S.R. (2021) The Pristimantis trachyblepharis species group, a clade of miniaturized frogs: description of four new species and insights into the evolution of body size in the genus. Zoological Journal of Linnean Society, 40, 1–40. https://doi.org/10.1093/zoolinnean/zlab044

  2.  

  3.  

  4.