Diversity, structure and natural history of amphibians in the upper Claro River basin, a buffer zone of the National Natural Park Los Nevados, Central Cordillera of Colombia

Main Article Content

Julián Andrés Rojas-Morales
Mateo Marín-Martínez


We present an assessment on composition, diversity and structure of amphibians in three zones along an elevation gradient (2,400–3,000 m) in the Central Cordillera of Colombia.  For this purpose, we carried out two field trips in November 2014 and February 2015, covering rainy and dry seasons, respectively.  Diurnal (08:00–12:00 h) and nocturnal (18:00–22:00 h) visual encounter surveys were made without spatial restrictions.  The diversity for each zone (alpha) and for the entire landscape (gamma) was evaluated by the effective number of species, and the structure of the communities was analyzed by range-abundance curves.  The inequality factor for each of the sampling zones was also calculated.  A total of 15 species belonging to seven genera and three families were recorded, all of the order Anura.  Craugastoridae with 11 species (73.3% of richness) and Pristimantis (eight species) were the most diverse family and genus, respectively.  The average alpha diversity per zone was 6.6 effective species, with zone A being the most diverse with eight species.  In terms of beta diversity we found 2.5 effective communities at the landscape level, and differences between zones are given by the rare species, while the most abundant ones (e.g., Pristimantis uranobates) are shared between them.  It is presumed that the greater diversity of zone A is due to the lower elevation and better state of conservation if compared to the other two zones.  Of the total species recorded, three are threatened with extinction: Endangered (Hypodactylus latens, Osornophryne percrassa), and Critically Endangered (Niceforonia adenobrachia).  The finding of three yet undescribed species is highlighted.

Article Details

How to Cite
Rojas-Morales, J.A. and Marín-Martínez, M. 2019. Diversity, structure and natural history of amphibians in the upper Claro River basin, a buffer zone of the National Natural Park Los Nevados, Central Cordillera of Colombia. Journal of Threatened Taxa. 11, 3 (Feb. 2019), 13261–13277. DOI:https://doi.org/10.11609/jott.4075.11.3.13261-13277.


Acevedo, A.A., R. Franco & D.A. Carrero (2016). Diversity of Andean amphibians of the Tamá National Natural Park in Colombia: a survey for the presence of Batrachochytrium dendrobatidis. Animal Biodiversity and Conservation 39(1): 1–10; https://abc.museucienciesjournals.cat/files/ABC_39-1_pp_1-10.pdf

Acosta, A.R. (2000). Ranas, Salamandras y Caecilias (Tetrapoda: Amphibia) de Colombia. Biota Colombiana 1(3): 289–319; https://redalyc.org/pdf/491/49110303.pdf

Acosta-Galvis, A.R. (2009). Estado del conocimiento de los anfibios y reptiles del departamento de Caldas. Vacíos de información y las prioridades de conservación. Corporación Autónoma Regional de Caldas (Corpocaldas). Manizales, Caldas. 142 pp.

Acosta, A.R. & D. Cuentas (2018). Lista de los Anfibios de Colombia: Referencia en línea V.05.2015.0 (acceso 02-04-2016). Página web accesible en https://www.batrachia.com; Batrachia, Villa de Leyva, Boyacá, Colombia.

Adams, W.M., R. Aveling, D. Brockington, B. Dickson, J. Elliot, J. Hutton, D. Roe, B. Vira & W. Womer (2004). Biodiversity conservation and the eradication of poverty. Science 306(5699): 1146–1149; https://www.researchgate.net/publication/51370569_Biodiversity_Conservation_and_the_Eradication_of_Poverty

Ardila-Robayo, M.C., P.M. Ruiz-Carranza & M. Barrera-Rodriguez (1996). Una nueva especie de Phrynopus (Amphibia, Anura, Leptodactylidae) de la Cordillera Central Colombiana. Lozania 67: 1–10.

Arroyo, S., A. Jerez & M.P. Ramírez-Pinilla (2003). Anuros de un bosque de niebla de la cordillera Oriental de Colombia. Caldasia 25(1): 153–167; https://www.unal.edu.co/icn/publicaciones/caldasia/25(1)/zoologia1.pdf

Arroyo, S.B., V.H. Serrano-Cardozo & M.P. Ramírez-Padilla (2008). Diet, microhabitat and time of activity in a Pristimantis (Anura, Strabomantidae) assemblage. Phyllomedusa 7(2): 109–119; https://doi.org/10.11606/issn.2316-9079.v7i2p109-119

Bernal, M.H. & J.D. Lynch (2008). Review and analysis of altitudinal distributions of the Andean anurans in Colombia. Zootaxa 1826: 1–25; https://www.researchgate.net/publication/276505871_Review_and_analysis_of_altitudinal_distribution_of_the_Andean_anurans_in_Colombia

Bolívar, W., L.A. Coloma, S. Ron & J.M. Rengifo (2010). Hyloscirtus larinopygion. The IUCN Red List of Threatened Species 2010: e.T55531A11328178. Downloaded on 17 March 2017; https://doi.org/10.2305/IUCN.UK.2010-2.RLTS.T55531A11328178.en

Cáceres-Andrade, S.P. & J.N. Urbina-Cardona (2009). Ensamblajes de anuros de sistemas productivos y bosques en el piedemonte llanero, departamento del Meta, Colombia. Caldasia 31(1): 175–194; https://revistas.unal.edu.co/index.php/cal/article/view/36085/37485

Cadavid, J.G., C. Román-Valencia & A.F. Gómez (2005). Composición y estructura de anfibios en un transecto altitudinal de los Andes centrales de Colombia. Revista del Museo Argentino de Ciencias Naturales 7(2): 103–118.

Casas-Pinilla, L.C., O. Mahecha-C, J.C. Dumar-R & C. Ríos-Maláver (2017). Diversidad de mariposas en un paisaje de bosque seco tropical, en la Mesa de los Santos, Santander, Colombia (Lepidoptera: Papilionoidea). SHILAP Revista de Lepidopterología 45(177): 83–108; https://www.redalyc.org/articulo.oa?id=45550375010

Castro, F., M.I. Herrera & J.D. Lynch (2004). Hypodactylus latens. The IUCN Red List of Threatened Species 2004: e.T56702A11517360. Downloaded on 07 January 2017; https://doi.org/10.2305/IUCN.UK.2004.RLTS.T56702A11517360

Chao, A. & L. Jost (2012). Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93(12): 2533–2547; https://doi.org/10.1890/11-1952.1

Cochran, D.M & J.C. Goin (1970). Frogs of Colombia. Publications of the United States National Museum. Smithsonian Institution 288: 1–641.

Cortés, C., A.M. Suárez-Mayorga& F.J. López-López (2006). Preparación y preservación de material científico, pp173–220. In: Angulo, A., J.V. Rueda-Almonacid, J.V. Rodríguez-Mahecha & E. La Marca (eds.). Técnicas de inventario y monitoreo para los anfibios de la región tropical andina. Conservación Internacional, Bogotá D.C., Colombia.

Dixo, M. & M. Martins (2008). Are leaf-litter frogs and lizards affected by edge effects due to forest fragmentation in Brazilian Atlantic forest? Journal of Tropical Ecology 24: 551–554; https://www.cambridge.org/core/journals/journal-of-tropical-ecology/article/are-leaflitter-frogs-and-lizards-affected-by-edge-effects-due-to-forest-fragmentation-in-brazilian-atlantic-forest/CD8F424D246CDDF9358D2BFE9C15D103

Dodson, C.H. & A.H. Gentry (1991). Biological extinction in western Ecuador. Annals of the Missouri Botanical Garden 78: 273–295.

Duellman, W.E. (1973). Descriptions of new Hylid frogs from Colombia and Ecuador. Herpetologica 29: 219–227; https://sci-hub.tw/https://www.jstor.org/stable/3892005

Duellman, W.E. & E. Lehr (2009). Terrestrial-Breeding Frogs (Strabomantidae) in Peru. Münster: Natur und Tier-Verlag GmbH. 382 p.

Duellman, W.E. & T.J. Berger (1982). A new species of Andean tree frog (Hylidae). Herpetologica 38: 456–460.

Duellman, W.E. & D.M. Hillis (1990). Systematic of frogs of the Hyla larinopygion group. Occasional Papers of the Museum of Natural History, The University of Kansas, Lawrence, Kansas 134: 1–23; https://archive.org/details/biostor-212

Ehrlich, P.R. & E.O. Wilson (1991). Biodiversity studies: Science and policy. Science 253(5021): 758–762.

Faivovich, J., C.F.B. Haddad, P.C.A. García, D.R. Frost & J.A. Campbell (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; https://digitallibrary.amnh.org/bitstream/handle/2246/462/B294.pdf?sequence=1

Frost, D.R. (2017). Amphibians Species of the World: an online reference. Version 6.0 (last Access on 25 September 2017). Electronic database accesible at https://research.amnh.org/herpetology/amphibia/index.html. American Museum of Natural History, New York, USA.

García J.C., H. Cárdenas & -F. Castro-H (2007). Relación entre la diversidad de anuros y los estados sucesionales de un bosque muy húmedo montano bajo del Valle del Cauca, suroccidente colombiano. Caldasia 29: 363–374; https://revistas.unal.edu.co/index.php/cal/article/view/39203/41056

Garcia, R.J.C., C.E. Posso-Gómez & H. Cárdenas-Henao (2015). Diet of Direct-Developing frogs (Anura: Craugastoridae: Pristimantis) from the Andes western Colombia. Acta Biológica Colombiana 20 (1): 79–87; https://www.scielo.org.co/pdf/abc/v20n1/v20n1a09.pdf

Greene, R.E., A. Balmford, P.R. Crane, G.M. Mace, J.D. Reynolds & R.K. Turner (2005). A framework for improved monitoring of biodiversity: Response to the world Summit on sustainable development. Conservation Biology 19(1): 56–65; https://johnreynolds.org/wp-content/uploads/2012/07/green-et-al-cons-bio-05.pdf

Gutiérrez-Cárdenas, P.D.A., K. Castillo, D. Martínez, C.F. Rocha & M.A. Rojas-Rivera (2016). Trophic ecology of Pristimantis labiosus (Anura: Craugastoridae) from Southwestern Colombia. North-Western Journal of Zoology 12(1): 102-109; https://www.researchgate.net/publication/304042766_Trophic_ecology_of_Pristimantis_labiosus_Anura_Craugastoridae_from_south-western_Colombia

Hammer, Ø., D.A.T. Harper & P.D. Ryan (2001). Past: Paleontological Statistics Software Package for education and data analysis. Paleontología Electrónica 4: 1–9.

Hill, M.O. (1973). Diversity and evenness: A unifying notation and its consequences. Ecology 54: 427–432; https://www.fcnym.unlp.edu.ar/catedras/ecocomunidades/Hill%20-%20Diversity%20and%20Evenness--%20A%20Unifying%20Notation%20and%20Its%20Consequences.pdf

Holdridge, L.R. (ed.) (1982). Ecología basada en zonas de vida. San José, Costa Rica. IICA. 215 pp.

Hutter, C.R., J.M. Guayasamin & J.J. Wiens (2013). Explaining Andean megadiversity: the evolutionary and ecological causes of glassfrog elevational richness patterns. Ecology Letters 16(9): 1135–1144; https://doi.org/10.1111/ele.12148

Hutter, C.R., S.M. Lambert & J.J. Wiens (2017). Rapid diversification and time explain Amphibian richness at different scales in the Tropical Andes, earth’s most biodiverse Hotspot. The American Naturalist 190(6): 828–843; https://www.wienslab.com/Publications_files/Hutter_et_al_AmNat_2017.pdf

Hsieh, T.C., K.H. Ma & A. Chao (2016). iNEXT: An R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods in Ecology and Evolution 7(12): 1451–1456; https://chao.stat.nthu.edu.tw/wordpress/paper/120_pdf_appendix.pdf

IUCN SSC Amphibian Specialist Group (2015). Osornophryne percrassa. The IUCN Red List of Threatened Species 2015: e.T54858A60783778. Downloaded on 02 March 2017. https://doi.org/10.2305/IUCN.UK.2015-2.RLTS.T54858A60783778.en

Jost, L. (2006). Entropy and diversity. Oikos 113(2): 363–375.

Jost, L. (2010). The relation between evenness and diversity. Diversity 2: 207–232; https://www.researchgate.net/publication/41667724_The_Relation_between_Evenness_and_Diversity

Kaplan, M. (1997). On the status of Hyla bogerti Cochran and Goin. Journal of Herpetology 31: 536–541.

Kattan, G.H., P. Franco, V. Rojas & G. Morales (2004). Biological diversification in a complex region: a spatial analysis of faunistic diversity and biogeography of the Andes of Colombia. Journal of Biogeography 31: 1829–1839; https://doi.org/10.1111/j.1365-2699.2004.01109.x/abstract

La Marca, E., K. Lips, S. Lötters, R. Puschendorf, R. Ibáñez, J.V. Rueda-Almonacid, , R. Schulte, C. Marty, F. Castro, J. Manzanilla-Puppo, J.E. García-Pérez, F. Bolaños, G. Chaves, J.A. Pounds, E. Toral & B.E. Young (2005). Catastrophic population decline and extinctions in Neotropical harlequin frogs (Bufonidae: Atelopus). Biotropica 37: 190–201; https://doi.org/10.1111/j.1744-7429.2005.00026.x

Lampo, M., A. Rodríguez-Contreras, E. La Marca & P. Daszak (2006). A chytridiomycosis epidemic and a severe dry season precede the disappearance of Atelopus species from the Venezuelan Andes. Herpetological Journal 16: 395–402; https://www.researchgate.net/publication/233616507_A_chytridiomycosis_epidemic_and_a_severe_dry_season_precede_the_disappearance_of_Atelopus_species_from_the_Venezuelan_Andes

Lips, K.R., D.E. Green & R. Papendick (2003). Chytridiomycosis in wild frogs from southern Costa Rica. Journal of Herpetology 37: 215–218; https://www.nwhc.usgs.gov/publications/documents/03KRL.JH01.pdf

Lips, K.R., Mendelson, J.R. III., A. Muñoz-Alonso, L. Canseco-Márquez & D.G. Mulcahy (2004). Amphibian population declines in montane southern Mexico: resurveys of historical localities. Biological Conservation 119: 555–564; https://www.sciencedirect.com/science/article/pii/S0006320704000217

Lynch, J.D. (1975). The identity of the frog Eleutherodactylus conspicillatus (Günther), with descriptions of two related species from northwestern South America (Amphibia, Leptodactylidae). Contributions in Science. Natural History Museum of Los Angeles County 272: 1–19.

Lynch, J.D. (1986). Origins of the high Andean herpetological fauna. Pp. 478–499. En: Vuilleumier, F. & Monasterio, M. (Eds.). High Altitude Tropical Biogeography. Oxford University Press. Oxford, UK.

Lynch, J.D. (1989). Intrageneric relationships of mainland Eleutherodactylus (Leptodactylidae). I. A review of the frogs assigned to the Eleutherodactylus discoidalis species group. Contributions in Biology and Geology. Milwaukee Public Museum 79: 1–25.

Lynch, J.D. (1991). A new diminutive Eleutherodactylus from the Cordillera Central of Colombia (Amphibia: Leptodactylidae). Journal of Herpetology 25(3): 344–352.

Lynch, J.D. (1999a). Ranas pequeñas, la geometría de la evolución, y la especiación en los Andes colombianos. Revista de la Academia de Ciencias Exactas, Físicas y Naturales 23(86): 143–159.

Lynch, J.D. (1999b). Lista anotada y clave para las ranas (Género Eleutherodactylus) chocoanas del Valle del Cauca y apuntes sobre las especies de la Cordillera Occidental adyacente. Caldasia 21(2): 184–202.

Lynch, J.D. & W.E. Duellman (1980). The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publication, University of Kansas Museum of Natural History 69: 1–86.

Lynch, J.D. & W.E. Duellman (1997). Frogs of the genus Eleutherodactylus in western Ecuador: systematic, ecology and biogeography. Museum of Natural History. University of Kansas. Special publication No 23. 236 pp.; https://www.biodiversitylibrary.org/bibliography/7951#/summary

Lynch, J.D. & J.V. Rueda-Almonacid (1997). Three new frogs Eleutherodactylus: Leptodactylidae) from cloud forest in eastern Departamento Caldas, Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales 21: 131–142.

Lynch, J.D. & A.M. Suárez-Mayorga (2002). Análisis biogeográfico de los anfibios paramunos. Caldasia 24(2): 471–480; https://www.bdigital.unal.edu.co/37602/1/39427-175445-1-PB.pdf

Lynch, J.D., P.M. Ruiz-Carranza & M.C. Ardila-Robayo (1994). The identities of the Colombian frogs confused with Eleutherodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae). Occasional Papers of the Museum of Natural History, University of Kansas 170: 1–42; https://www.biodiversitylibrary.org/page/4466887#page/5/mode/1up

Lynch, J.D., P.M. Ruiz-Carranza & M.C. Ardila-Robayo (1996). Three new species of Eleutherodactylus (Amphibia: Leptodactylidae) from high elevations of the Cordillera Central of Colombia. Caldasia 18: 329–342.

Lynch, J.D., P.M. Ruiz-Carranza & M.C. Ardila-Robayo (1997). Biogeographic patterns of Colombians frogs and toads. Revista de la Academia de Ciencias Exactas, Físicas y Naturales 21: 237–248; https://zoologia.puce.edu.ec/vertebrados/Recursos/publicaciones/Cientifica/Lynch_and_Ruiz_Carranza1997.pdf

Mantilla-Castaño, J.C., S. Escobar-Lasso, J.A. Rojas-Morales & J.V. González-Arenas (in press). First record of the Colombian endemic toad Osornophryne percrassa Ruíz-Carranza and Hernández-Camacho 1976 (Anura: Bufonidae) from the department of Risaralda, Central Cordillera of Colombia. Revista de Biodiversidad Neotropical

Márquez, G. (2001). De la abundancia a la escasez: La transformación de ecosistemas en Colombia. Pp. 325–452. En: Palacio, G. (Ed.). Naturaleza en disputa. Ensayos de historia ambiental en Colombia 1850-1995. Universidad Nacional de Colombia, Bogotá, Colombia.

Méndez-Narváez, J. & W. Bolívar-G (2016). Complementary ecological approaches to understand decreases in frog diversity in altered Andean ecosystems. South American Journal of Herpetology 11(1): 1–11; https://www.researchgate.net/publication/301732403_Complementary_Ecological_Approaches_to_Understand_Decreases_in_Frog_Diversity_in_Altered_Andean_Ecosystems

Meza-Joya, F.L. & M. Torres (2016). Spatial diversity patterns of Pristimantis frogs in the Tropical Andes. Ecology and Evolution 6(7): 1901–1913; https://doi.org/10.1002/ece3.1968

Moen, D.S., S.A. Smith & J.J. Wiens. (2009). Community assembly through evolutionary diversification and dispersal in Middle American treefrogs. Evolution 63: 3228–3247; https://www.researchgate.net/publication/26725896_Community_Assembly_Through_Evolutionary_Diversification_and_Dispersal_in_Middle_American_Treefrogs

Myers, N., R.A. Mittermeier, C.G. Mittermeier, G.A.B. Da Fonseca & J. Kent (2000). Biodiversity hotspots for conservation priorities. Nature 403: 853–858; https://www.nature.com/articles/35002501

Navas, C.A. (2006). Patterns of distribution of anurans in high Andean tropical elevations: Insights from integrating biogeography and evolutionary physiology. Integrative and Comparative Biology 46(1): 82–91; https://www.researchgate.net/publication/51218756_Patterns_of_distribution_of_anurans_in_high_Andean_tropical_elevations_Insights_from_integrating_biogeography_and_evolutionary_physiology

Osorno, M., D.L. Gutiérrez-Lamus & J.C. Blanco (2011). Anfibios en un gradiente de intervención en el noroccidente de la Amazonia Colombiana. Revista Colombia Amazónica 4: 143–160; https://www.sinchi.org.co/revista-colombia-amazonica-iv

Ricotta, C. (2003). Parametric scaling from species relative abundances to absolute abundances in the computation of biological diversity: a first proposal using Shannon’s entropy. Acta Biotheoretica 51: 181–188; https://link.springer.com/article/10.1023/A%3A1025142106292?no-access=true

Rivera-Correa, M. & J. Faivovich (2013). A new species of Hyloscirtus (Anura: Hylidae) from Colombia, with a rediagnosis of Hyloscirtus larinopygion (Duellman, 1973). Herpetologica 69(3): 298–313; https://www.researchgate.net/publication/256458524_A_New_Species_of_Hyloscirtus_Anura_Hylidae_from_Colombia_with_a_Rediagnosis_of_Hyloscirtus_larinopygion_Duellman_1973

Rojas-Morales, J.A., H.F. Arias-Monsalve & G.A. González-Durán (2014a). Anfibios y reptiles de la región centro-sur del departamento de Caldas, Colombia. Biota Colombiana 15(1): 73–93; https://www.redalyc.org/pdf/491/49140738005.pdf

Rojas-Morales, J.A., A. Castro-Gómez & S. Escobar-Lasso (2014b). Joyas de los Andes Tropicales: Colombia, tierra de ranas de cristal. Bioma 19: 6–39; https://www.academia.edu/6948400/Joyas_de_los_Andes_tropicales_Colombia_tierra_de_ranas_de_cristal

Rös, M., F. Escobar & G. Halffter (2012). How dung beetles respond to a human-modified variegated landscape in Mexican cloud forest: a study of biodiversity integrating ecological and biogeographical perspectives. Diversity and Distributions 18(4): 377–389; https://www.researchgate.net/publication/230540150_How_dung_beetles_respond_to_a_human-modified_variegated_landscape_in_Mexican_cloud_forest_A_study_of_biodiversity_integrating_ecological_and_biogeographical_perspectives

Ruiz-Carranza, P.M. & J.I. Hernández-Camacho (1976). Osornophryne, género nuevo de anfibios bufónidos de Colombia y Ecuador. Caldasia 6: 93−148; https://revistas.unal.edu.co/index.php/cal/article/view/34333/34534

Santori, C. & N. McManus (2014). Herpetofaunal abundance and species diversity in coastal Ecuador: differences between forest types, the role of abiotic factors and edge effects. Tropical Conservation, Permaculture & Research Internship Jama-Coaque Ecological Reserve. 23 pp.

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

Urbina, C.J.N. & M.C. Londoño (2003). Distribución de la comunidad de herpetofauna asociada a cuatro áreas con diferente grado de perturbación en la isla Gorgona, pacífico colombiano. Revista de la Academia de Ciencias Exactas, Físicas y Naturales 27(102): 105–113; https://www.researchgate.net/profile/Nicolas_Urbina-Cardona/publication/216019359_Distribucion_de_la_comunidad_de_herpetofauna_asociada_a_cuatro_areas_con_diferente_grado_de_perturbacion_en_la_Isla_Gorgona_Pacifico_colombiano/links/07ab43354ba7e91da7405dc5/Distribucion-de-la-comunidad-de-herpetofauna-asociada-a-cuatro-areas-con-diferente-grado-de-perturbacion-en-la-Isla-Gorgona-Pacifico-colombiano.pdf

Valencia, A. (1985). La colonización y el desarrollo económico-social del Gran Caldas (Siglo XIX). Revista Universidad de Caldas 5(2): 146.

Vanegas-Guerrero, J. & J.D. Fernández-Roldán (2014). Osornophryne percrassa Ruiz-Carranza y Hernández-Camacho 1976. Catálogo de Anfibios y Reptiles de Colombia 2: 21−24.

Vanegas-Guerrero, J., G.A. González-Duran & S. Escobar-Lasso (2016). Distribution, diet, and vocalizations of the endangered Colombian toad Osornophryne percrassa (Anura, Bufonidae). Herpetological Conservation and Biology 11(1): 90−100; https://www.herpconbio.org/Volume_11/Issue_1/Vanegas-Guerrero_etal_2016.pdf

Vargas-Salinas, F. & A. Aponte-Gutiérrez (2016). Diversidad y recambio de anfibios y reptiles entre coberturas vegetales en una localidad del valle del Magdalena medio, departamento de Antioquia, Colombia. Biota Colombiana 17(2): 117−137; www.redalyc.org/pdf/491/49151352010.pdf

Vonesh, J.R. (2001). Patterns of richness and abundance in a tropical African leaf-litter herpetofauna. Biotropica 33(3): 502–510; https://doi.org/10.1111/j.1744-7429.2001.tb00204.x