ACOUSTIC VARIATION OF THE SAVANNAH FROG (Dendropsophus molitor) IN AN ELEVATION GRADIENT IN THE EAST COLOMBIAN MOUNTAIN RANGE

Laura Sofia Delgado Montufar; Angela M. Mendoza-Henao; Nelsy R. Pinto-Sánchez

doi:10.22201/fc.25942158e.2026.2.1410

Autores/as

Laura Sofia Delgado Montufar https://orcid.org/0009-0009-2025-8395
Angela M. Mendoza-Henao Red de Ecoacústica Colombiana https://orcid.org/0000-0002-6752-9238
Nelsy R. Pinto-Sánchez Universidad Militar Nueva Granada https://orcid.org/0000-0001-7561-2982

DOI:

https://doi.org/10.22201/fc.25942158e.2026.2.1410

Palabras clave:

Anura, adaptación, cantos de advertencia, tamaño corporal

Resumen

La variación de los cantos de advertencia puede llevar al aislamiento reproductivo cuando las parejas no reconocen dichos cantos. Por lo tanto, sistemas de alta montaña y las variaciones ambientales dentro de estos pueden moldear rasgos implicados en el reconocimiento de pareja y fomentar mecanismos de especiación. Dendropsophus molitor es un anuro endémico de los Andes que se distribuye en un amplio rango de elevación (2100–3500 m s.n.m.). Esta especie produce cantos de advertencia que pueden covariar con su temperatura corporal, tamaño corporal y la humedad relativa, factores que a su vez varían con la elevación en sistemas montañosos como los Andes. Para determinar si existe variación geográfica de los cantos de D. molitor en un gradiente de elevación, analizamos la relación entre la humedad relativa, temperatura corporal, y tamaño corporal a lo largo de un gradiente de elevación. Posteriormente, utilizamos modelos lineales mixtos para evaluar la relación entre las variables de respuesta relacionadas con los cantos (incluyendo los parámetros temporales y espectrales) y las variables predictoras (efectos fijos) ambientales (humedad relativa y temperatura corporal) y morfológicas. Observamos que la temperatura y tamaño corporal, pero no la humedad relativa, cambian significativamente a lo largo del gradiente de elevación. Así mismo detectamos una variación geográfica en la mayoría de los parámetros de los cantos de D. molitor, explicada principalmente por el tamaño corporal. Estos resultados sugieren que los gradientes de elevación en sistemas montañosos pueden moldear indirectamente rasgos relacionados con la reproducción y potencialmente contribuir al aislamiento reproductivo.

Citas

Adams, D. C., & Church, J. O. (2008). Amphibians do not follow Bergman’s rule. Evolution, 63(1), 413–420. https://doi.org/10.1111/j.1558-5646.2007.00297.x

Altunisik, A., & Ozdemir, N. (2013). Body size and age structure of a highland population of Hyla orientalis Bedriaga, 1890, in northern Turkey. Herpetozoa, 26(1-2), 49–55.

Angilletta, M. J., Niewiarowski, P. H., & Navas, C. A. (2002). The evolution of thermal physiology in ectotherms. Journal of Thermal Biology, 27(4), 249–268. https://doi.org/10.1016/S0306-4565(01)00094-8

Annibale, F. S., de Sousa, V. T. T., da Silva, F. R., & Murphy, C. G. (2020). Geographic Variation in the Acoustic Signals of Dendropsophus nanus (Boulenger 1889) (Anura: Hylidae). Herpetologica, 76(3), 267–277. https://doi.org/10.1655/Herpetologica-D-19-00046.1

Araya-Salas, M., & Smith-Vidaurre, G. (2017). WarbleR: An R package to streamline analysis of animal acoustic signals. Methods in Ecology and Evolution, 8(2), 184–191. https://doi.org/10.1111/2041-210X.12624

Arias-Cárdenas, A., Barrientos, L. S., Pardo-Diaz, C., Paz, A., Crawford, A. J., & Salazar, C. (2024). Taxonomic inflation and a reconsideration of speciation in the Andes: The case of the high-elevation tree frog Dendropsophus molitor (Anura: Hylidae). Zoological Journal of the Linnean Society, 200(3), 763–775. https://doi.org/10.1093/zoolinnean/zlad085

Ashton, K. G. (2002). Do amphibians follow Bergmann’s rule? Canadian Journal of Zoology, 80(4), 708–716. https://doi.org/10.1139/z02-049

Bell, P. L. (2012). Geografía, topografía y clima. En: Banco de la República (Ed.), Colombia: Manual comercial e industrial (pp. 37–50). Banco de la República de Colombia.

Bader, M. Y., Rietkerk, M., & Bregt, A. K. (2007). Vegetation Structure and Temperature Regimes of Tropical Alpine Treelines. Arctic, Antarctic, and Alpine Research, 39(3), 353–364. http://www.jstor.org/stable/20181707

Bates, D., Mächler, M., Bolker, B., & Walker, S. (2015). Fitting linear mixed-effects models using lme4. Journal of Statistical Software, 67(1), 1–48. https://doi.org/10.18637/jss.v067.i01

Bernal, X. E., Guarnizo, C., & Lüddecke, H. (2005). Geographic variation in advertisement call and genetic structure of Colostethus palmatus (anura, Dendrobatidae) from the Colombian Andes. Herpetologica, 61(4), 395–408. https://doi.org/10.1655/04-87.1

Bosch, J., & De la Riva, I. (2004). Are frog calls modulated by the environment? An analysis with anuran species from Bolivia. Canadian Journal of Zoology, 82(6), 880–888. https://doi.org/10.1139/z04-060

Boschetti, J. P., De Bastiani, V. I. M., Lingnau, R., & Lucas, E. M. (2019). Bioacoustics of Pithecopus rusticus (Anura, Phyllomedusidae): A Rare Species Possibly Threatened with Extinction. South American Journal of Herpetology, 14(3), 196–203. https://doi.org/10.2994/SAJH-D-17-00071.1

Boschman, L. M., & Condamine, F. L. (2022). Mountain radiations are not only rapid and recent: Ancient diversification of South American frog and lizard families related to Paleogene Andean orogeny and Cenozoic climate variations. Global and Planetary Change, 208, 103704. https://doi.org/10.1016/j.gloplacha.2021.103704

Campos-Cerqueira, M., & Aide, T. M. (2017). Changes in the acoustic structure and composition along a tropical elevational gradient. Journal of Ecoacoustics, 1, 1–13. https://doi.org/10.22261/JEA.PNCO7I

Caro, L. M., Caycedo-Rosales, P. C., Bowie, R. C. K., Slabbekoorn, H., & Cadena, C. D. (2013). Ecological speciation along an elevational gradient in a tropical passerine bird? Journal of Evolutionary Biology, 26(2), 357–374. https://doi.org/10.1111/jeb.12055

Colombini, B., Nocella, M., Benelli, G., Cecchi, G., & Bagni, M. A. (2008). Effect of temperature on cross-bridge properties in intact frog muscle fibers. American Journal of Physiology-Cell Physiology, 294(4), 1113–1117. https://doi.org/10.1152/ajpcell.00063.2008

Cortés, J. E. (2009). Requerimientos de hábitat en un ensamblaje de anuros en dos tipos de cobertura, en el municipio de Villa de Leyva (Departamento de Boyacá) [Bachelor's thesis, Pontificia Universidad Javeriana]. http://repository.javeriana.edu.co/handle/10554/14567

Cui, J. G., Song, X. Y., Fang, G. Z., Brauth, S. E., Xu, F., & Tang, Y. Z. (2011). Circadian Rhythm of Calling Behavior in the Emei Music Frog (Babina daunchina) is Associated with Habitat Temperature and Relative Humidity. Asian Herpetological Research, 2(3), 1490–154. https://doi.org/10.3724/SP.J.1245.2011.00149

Delignette-Muller, M. L., & Dutang, C. (2015). fitdistrplus: An R package for fitting distributions. Journal of Statistical Software, 64(4), 1–34. https://doi.org/10.18637/jss.v064.i04

De Toledo, L. F., Fernado, C., & Haddad, B. (2009). Defensive vocalizations of neotropical anurans. Journal of herpetology, 4(1), 25–42. https://doi.org/doi:10.2994/057.004.0104

Esquerre, D., Brennan, I., Catullo, R., Torres-Perez, F., & Keogh, J. (2018). How mountains shape biodiversity: The role of the Andes in biogeography, diversification, and reproductive biology in South America’s most species-rich lizard radiation (Squamata: Liolaemidae). Evolution, 73(2), 214–230. https://doi.org/10.1111/evo.13657

Fang, G., Jiang, F., Yang, P., Cui, J., Brauth, S. E., & Tang, Y. (2014). Male vocal competition is dynamic and strongly affected by social contexts in music frogs. Animal Cognition, 17(2), 483–494. https://doi.org/10.1007/s10071-013-0680-5

Gerhardt, H. C. (1991). Female mate choice in treefrogs: Static and dynamic acoustic criteria. Animal Behaviour, 42(4), 615–635. https://doi.org/10.1016/S0003-3472(05)80245-3

Gerhardt, H. C., & Huber, F. (2002). Acoustic Communication in Insects and Anurans: Common Problems and Diverse Solutions. University of Chicago Press.

Gheyret, G., Mohammat, A., & Tang, Z. (2020). Elevational patterns of temperature and humidity in the middle Tianshan Mountain area in Central Asia. Journal of Mountain Science, 17(2), 397–409. https://doi.org/10.1007/s11629-019-5481-0

Gillooly, J. F., & Ophir, A. G. (2010). The energetic basis of acoustic communication. Proceedings of the Royal Society B: Biological Sciences, 277(1686), 1325–1331. https://doi.org/10.1098/rspb.2009.2134

Gingras, B., Boeckle, M., Herbst, C. T., & Fitch, W. T. (2013). Call acoustics reflect body size across four clades of anurans. Journal of Zoology, 289(2), 143–150. https://doi.org/10.1111/j.1469-7998.2012.00973.x

Goicoechea, N., De La Riva, I., & Padial, J. (2010). Recovering phylogenetic signal from frog mating calls. Zoologica Scripta, 39, 411–154. https://doi.org/10.1111/j.1463-6409.2009.00413.x

Gridi-Papp, M. (2014). Is the frequency content of the calls in North American treefrogs limited by their larynges? International Journal of Evolutionary Biology, 2014(1), 198069. https://doi.org/10.1155/2014/198069

Guarnizo, C., Armesto, L. O., & Acevedo, A. (2014). Dendropsophus labialis (Peters, 1863): Rana Andina. Catálogo de Anfibios y Reptiles de Colombia, 2 (2), 56–71.

Guarnizo, C. E., Amézquita, A., & Bermingham, E. (2009). The relative roles of vicariance versus elevational gradients in the genetic differentiation of the high Andean tree frog, Dendropsophus labialis. Molecular Phylogenetics and Evolution, 50(1), 84–92. https://doi.org/10.1016/j.ympev.2008.10.005

Guarnizo, C. E., Escallón, C., Cannatella, D., & Amézquita, A. (2012). Congruence between acoustic traits and genealogical history reveals a new species of Dendropsophus (Anura: Hylidae) in the High Andes of Colombia. Herpetologica, 68(4), 523–540. https://doi.org/10.1655/HERPETOLOGICA-D-10-00038

Gutiérrez, G., & Lüddecke, H. (2002). Mating pattern and hatching success in a population of the Andean frog Hyla labialis. Amphibia-Reptilia, 23(3), 281–292. https://doi.org/10.1163/15685380260449162

Harris, C. M. (1966). Absorption of sound in air versus humidity and temperature. The Journal of the Acoustical Society of America, 40(1), 148–159. https://doi.org/10.1121/1.1910031

Higuera-Rojas, D. F., & Carvajal-Cogollo, J. E. (2021). Diet of Dendropsophus molitor (Anura: Hylidae) in a High-Andean agricultural ecosystem, Colombia. Universitas Scientiarum, 26(1), 119–137. https://doi.org/10.11144/Javeriana.SC26-1.dodm

Hoorn, C., Wesselingh, F. P., Ter Steege, H., Bermudez, M., Mora, A., Sevink, J., & Antonelli, A. (2010). Amazonia through time: andean uplift, climate change, landscape evolution, and biodiversity. Science, 330(6006), 927–931. https://www.science.org/doi/10.1126/science.1194585

Hopkins, G. R., & Lahanas, P. N. (2011). Aggregation behavior in a neotropical dendrobatid frog (Allobates talamancae) in western Panama. Behaviour, 148(3), 359–372. http://www.jstor.org/stable/23034198

Hoskin, C. J., James, S., & Grigg, G. C. (2009). Ecology and taxonomy-driven deviations in the frog call–body size relationship across the diverse Australian frog fauna. Journal of Zoology, 278(1), 36–41. https://doi.org/10.1111/j.1469-7998.2009.00550.x

Hutchison, V. H., Whitford, W. G., & Kohl, M. (1968). Relation of body size and surface area to gas exchange in anurans. Physiological Zoology, 41(1), 65–85. http://www.jstor.org/stable/30158485

Khatiwada, J. R., Zhao, T., & Jiang, J. (2020). Variation of body temperature of active amphibians along elevation gradients in eastern Nepal Himalaya. Journal of Thermal Biology, 92, 102653. https://doi.org/10.1016/j.jtherbio.2020.102653

Köhler, A., Sadowska, J., Olszewska, J., Trzeciak, P., Berger-Tal, O., & Tracy, C. R. (2011). Staying warm or moist? Operative temperature and thermal preferences of common frogs (Rana temporaria), and effects on locomotion. Herpetological Journal, 21(1), 17–26.

Köhler, J., Jansen, M., Rodríguez, A., Kok, P. J. R., Toledo, L. F., Emmrich, M., Glaw, F., Haddad, C. F. B., Rödel, 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.4251.1.1

Lardner, B., & Lakim, M. B. (2004). Female call preferences in tree-hole frogs: Why are there so many unattractive males? Animal Behaviour, 68(2), 265–272. https://doi.org/10.1016/j.anbehav.2004.05.003

Larter, L. C., & Ryan, M. C. (2024). Túngara frog call-timing decisions arise as internal rhythms interact with fluctuating chorus noise. Behavioral Ecology, 35(4), 1–14. https://doi.org/10.1093/beheco/arae034

Laugen, A. T., Laurila, A., Jönsson, K. I., Söderman, F., & Merilä, J. (2005). Do common frogs (Rana temporaria) follow Bergmann’s rule? Evolutionary Ecology Research, 717–731.

Loman, J. (2002). Temperature, genetic and hydroperiod effects on metamorphosis of brown frogs Rana arvalis and R. temporaria in the field. Journal of Zoology, 258(1), 115–129. https://doi.org/10.1017/S0952836902001255

Lomolino, M. (2001). Elevation gradients of species density: Historical and prospective views. Global Ecology and Biogeography, 10(1), 3–13. https://doi.org/doi:10.1046/j.1466-822x.2001.00229.x

Lüddecke, H., Amézquita, A., Bernal, X., & Guzmán, F. (2000). Partitioning of vocal activity in a Neotropical highland-frog community. Studies on Neotropical Fauna and Environment, 35(3), 185–194. https://doi.org/10.1076/snfe.35.3.185.8855

Lüddecke, H., & Sánchez, O. R. (2002). Are tropical highland frog calls cold-adapted? The case of the Andean Frog Hyla labialis. Biotropica, 34(2), 281–288.

Ma, X., Tong, L., & Lu, X. (2009). Variation of body size, age structure and growth of a temperate frog, Rana chensinensis, over an elevational gradient in Northern China. Amphibia-Reptilia, 30, 11–17. https://doi.org/10.1163/156853809787392685

Martin, W. F. (1971). Mechanics of sound production in toads of the genus Bufo: Passive elements. Journal of Experimental Zoology, 176(3), 273–293. https://doi.org/10.1002/jez.1401760304

Martínez Baños, V., Pacheco Florez, V., & Ramírez-Pinilla, M. P. (2011). Relative abundance and microhabitat use by the frog Geobatrachus walkeri (Anura: Strabomantidae) in two habitats of Sierra Nevada de Santa Marta, Colombia. Revista de biologia tropical, 59(2), 907–920.

Mendoza-Henao, A. M., Zamudio, K. R., Guayasamin, J. M., Escalona, M., & Parra-Olea, G. (2023). Environment rather than character displacement explains call evolution in glassfrogs. Evolution, 77(2), 355–369. https://doi.org/10.1093/evolut/qpac041

Morais, A. R., Batista, V. G., Gambale, P. G., Signorelli, L., & Bastos, R. P. (2012). Acoustic communication in a Neotropical frog (Dendropsophus minutus): Vocal repertoire, variability, and individual discrimination. The Herpetological Journal, 22(4), 249–257.

Moreno-Gómez, F. N., Bacigalupe, L. D., Silva-Escobar, A. A., & Soto-Gamboa, M. (2015). Female and male phonotactic responses and the potential effect of sexual selection on the advertisement calls of a frog. Animal Behaviour, 104, 79–86. https://doi.org/10.1016/j.anbehav.2015.03.010

Moser, C. F., Schuck, L. K., Olmedo, G. M., & Lingnau, R. (2022). Individual variation in the advertisement call of Aplastodiscus albosignatus (Anura: Hylidae) is correlated with body size and environmental temperature. Zoologia (Curitiba), 39, e21008. https://doi.org/10.1590/S1984-4689.v39.e21008

Motegi, T., Mizutani, K., & Wakatsuki, N. (2013). Simultaneous measurement of air temperature and humidity based on sound velocity and attenuation using ultrasonic probe. Japanese Journal of Applied Physics, 52(7S), 07HC05. https://doi.org/10.7567/JJAP.52.07HC05

Nali, R. C., & Prado, C. P. (2014). The fight call of Bokermannohyla ibitiguara (Anura: Hylidae): First record for the genus. Salamandra, 50, 181–184.

Narins, P., & Meenderink, S. (2014). Climate change and frog calls: Long-term correlations along a tropical altitudinal gradient. Proceedings of the Royal Society B: Biological Sciences, 281(1783), 20140401. https://doi.org/10.1098/rspb.2014.0401

Navas, C. A. (1996a). Implications of microhabitat selection and patterns of activity on the thermal ecology of high elevation neotropical anurans. Oecologia, 108(4), 617–626. https://doi.org/10.1007/BF00329034

Navas, C. A. (1996b). Thermal dependency of field locomotor and vocal performance of high-elevation anurans in the Tropical Andes. Journal of Herpetology, 30(4), 478–487. https://doi.org/10.2307/1565690

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://doi.org/10.1093/icb/icj001

Navas, C. A., & Bevier, C. R. (2001). Thermal dependency of calling performance in the eurythermic frog Colostethus subpunctatus. Herpetologica, 57(3), 384–395. https://www.jstor.org/stable/3893104

Navas, C. A., Carvajalino-Fernández, J. M., Saboyá-Acosta, L. P., Rueda-Solano, L. A., & Carvajalino-Fernández, M. A. (2013). The body temperature of active amphibians along a tropical elevation gradient: Patterns of mean and variance and inference from environmental data. Functional Ecology, 27(5), 1145–1154. https://doi.org/10.1111/1365-2435.12106

Navas, C. A., Gomes, F. R., & Carvalho, J. E. (2008). Thermal relationships and exercise physiology in anuran amphibians: Integration and evolutionary implications. Comparative Biochemistry and Physiology Part A: Molecular & Integrative Physiology, 151(3), 344–362. https://doi.org/10.1016/j.cbpa.2007.07.003

Olalla-Tárraga, M. Á., & Rodríguez, M. Á. (2007). Energy and interspecific body size patterns of amphibian faunas in Europe and North America: Anurans follow Bergmann’s rule, urodeles its converse. Global Ecology and Biogeography, 16(5), 606–617. https://doi.org/10.1111/j.1466-8238.2007.00309.x

Ortiz, E., & Bejarano, L. (2023). El estudio del desarrollo embrionario en Dendropsophus molitor como problema de conocimiento [Master thesis], Universidad Pedagógica Nacional.

Ota, K., Aihara, I., & Aoyagi, T. (2020). Interaction mechanisms quantified from dynamical features of frog choruses. Royal Society Open Science, 7(3), 191693. https://doi.org/10.1098/rsos.191693

Rangwala, I., & Miller, J. R. (2012). Climate change in mountains: A review of elevation-dependent warming and its possible causes. Climatic Change, 114(3), 527–547. https://doi.org/10.1007/s10584-012-0419-3

R Core Team. (2024). R: A language and environment for statistical computing. R Foundation for Statistical Computing. https://www.R-project.org/

Reichert, M. S., & Gerhardt, H. C. (2011). The role of body size on the outcome, escalation, and duration of contests in the grey treefrog, Hyla versicolor. Animal Behaviour, 82(6), 1357-1366. https://doi.org/10.1016/j.anbehav.2011.09.019

Rodríguez, R. L., Araya-Salas, M., Gray, D. A., Reichert, M. S., Symes, L. B., Wilkins, M. R., Safran, R. J., & Höbel, G. (2015). How acoustic signals scale with individual body size: Common trends across diverse taxa. Behavioral Ecology, 26(1), 168–177. https://doi.org/10.1093/beheco/aru174

Rozen-Rechels, D., Dupoué, A., Lourdais, O., Chamaillé-Jammes, S., Meylan, S., Clobert, J., & Le Galliard, J.-F. (2019). When water interacts with temperature: Ecological and evolutionary implications of thermo-hydroregulation in terrestrial ectotherms. Ecology and Evolution, 9(17), 10029–10043. https://doi.org/10.1002/ece3.5440

Sanín Montoya, A. (2003). La honestidad en las llamadas agresivas de Hyla Labialis (Anura: Hylidae) [Bachelor's thesis], Universidad de los Andes.

Smith, B. T., McCormack, J. E., Cuervo, A. M., Hickerson, M. J., Aleixo, A., Cadena, C. D., Pérez-Emán, J., Burney, C. W., Xie, X., Harvey, M. G., Faircloth, B. C., Glenn, T. C., Derryberry, E. P., Prejean, J., Fields, S., & Brumfield, R. T. (2014). The drivers of tropical speciation. Nature, 515(7527), 406–409. https://doi.org/10.1038/nature13687

Sueur, J., Aubin, T., & Simonis, C. (2008). Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18(2), 213–226. https://doi.org/10.1080/09524622.2008.9753600

Tessarolo, G., Maciel, N. M., Ribeiro Morais, A., & Bastos, R. P. (2016). Geographic variation in advertisement calls among populations of Dendropsophus cruzi (Anura: Hylidae). The Herpetological Journal, 26(3), 219–227. https://www.jstor.org/stable/27098788

Tonini, J. F. R., Provete, D. B., Maciel, N. M., Morais, A. R., Goutte, S., Toledo, L. F., & Pyron, R. A. (2020). Allometric escape from acoustic constraints is rare for frog calls. Ecology and Evolution, 10(8), 3686–3695. https://doi.org/10.1002/ece3.6155

Turin, R. A. F., Nali, R. C., & Prado, C. P. A. (2018). Intraspecific call variation in a Neotropical gladiator frog with a complex advertisement call. Amphibia-Reptilia, 39(1), 31–39. https://doi.org/10.1163/15685381-00003147

Valdivieso, D., & Tamsitt, J. R. (1974). Thermal relations of the neotropical frog, Hyla labialis (Anura: Hylidae). Royal Ontario Museum; Life science occasional papers, 26, 1–12.

Valetti, J. A., & Martino, A. L. (2012). Temperature effect on the advertisement call of Pleurodema tucumanum (Anura: Leiuperidae). Phyllomedusa Journal of Herpetology, 11(2), 125–134. https://doi.org/10.11606/issn.2316-9079.v11i2p125-134

Veiga-Chaves, M. V., Alcântara de Sena, V. M. A., & Diniz, P. (2024). Geographic variation in the acoustic signals of lesser treefrogs Dendropsophus minutus (Anura, Hylidae). Papéis Avulsos de Zoologia, 64, e202464017. https://doi.org/10.11606/1807-0205/2024.64.017

Vilaça, T. R. A., Silva., Rodrigues dos Santos, J., & and Solé, M. (2011). Vocalization and territorial behaviour of Phyllomedusa nordestina Caramaschi, 2006 (Anura: Hylidae) from southern Bahia, Brazil. Journal of Natural History, 45(29-30), 1823–1834. https://doi.org/10.1080/00222933.2011.561018

Watters, J. L., Cummings, S. T., Flanagan, R. L., & Siler, C. D. (2016). Review of morphometric measurements used in anuran species descriptions and recommendations for a standardized approach. Zootaxa, 4072(4), 477–495. https://doi.org/10.11646/zootaxa.4072.4.6

Wei, T., & Simko, V. (2021). R package 'corrplot': Visualization of a correlation matrix (Version 0.92) [R package]. Comprehensive R Archive Network. https://github.com/taiyun/corrplot

Wells, K. D. (2010). The Ecology and Behavior of Amphibians. University of Chicago Press.

Wickham, H., François, R., Henry, L., Müller, K., & Vaughan, D. (2023). dplyr: A grammar of data manipulation (Version 1.1.3) [R package]. Comprehensive R Archive Network. https://CRAN.R-project.org/package=dplyr

Wilczynski, W., McClelland, B. E., & Rand, A. S. (1993). Acoustic, auditory, and morphological divergence in three species of neotropical frog. Journal of Comparative Physiology A, 172(4), 425–438. https://doi.org/10.1007/BF00213524

Zalakeviciute, R., López-Villada, J., & Rybarczyk, Y. (2018). Contrasted effects of relative humidity and precipitation on urban PM2.5 pollution in high elevation urban areas. Sustainability, 10(6), 2064. https://doi.org/10.3390/su10062064

Ziegler, L., Arim, M., & Bozinovic, F. (2016). Intraspecific scaling in frog calls: The interplay of temperature, body size and metabolic condition. Oecologia, 181(3), 673–681. https://doi.org/10.1007/s00442-015-3499-8

Zhao, L., Santos, J.C., Wang, J., Ran, J., Tang, Y., & Cui, J. Noise constrains the evolution of call frequency contours in flowing water frogs: a comparative analysis in two clades. Frontiers in Zoology, 18, 37. https://doi.org/10.1186/s12983-021-00423-y

Zsemberovszky, L. (2018). Vocal behavior and intraspecific call variations of Kempholey Night frog (Nyctibatrachus kempholeyensis) in the Western Ghats, India [master’s thesis], Norwegian University of Life Sciences.