Llamadas repetitivas en la Ratona Común Sudamericana: la información codificada en el número y tasa de repetición de los llamados de alarma

Gustavo J. Fernández; Mariana E. Carro

doi:10.56178/eh.v41i2.1543

Artículos Publicado 24 septiembre 2026 https://doi.org/10.56178/eh.v41i2.1543

Llamadas repetitivas en la Ratona Común Sudamericana: la información codificada en el número y tasa de repetición de los llamados de alarma

comunicación llamadas de disuasión tasas de repetición Troglodytes musculus

Gustavo J. Fernández Grupo de Investigación en Ecofisiología de Fauna Silvestre, INIBIOMA-CONICET, Centro Regional Universitario San Martín de los Andes (CRUSMA). Universidad Nacional del Comahue, Pasaje de la Paz 235, (8370) San Martín de los Andes, Neuquén, Argentina

Mariana E. Carro Grupo de Investigación en Ecofisiología de Fauna Silvestre, INIBIOMA-CONICET, Centro Regional Universitario San Martín de los Andes (CRUSMA). Universidad Nacional del Comahue, Pasaje de la Paz 235, (8370) San Martín de los Andes, Neuquén, Argentina

Resumen

Las aves frecuentemente utilizan señales acústicas de alarma para alertar a co- y heterospecíficos de la presencia cercana de una amenaza. Los emisores pueden usar una señal única o largas secuencias de llamados repetitivos las cuales pueden producir un efecto sostenido en el comportamiento de los receptores de la señal. La tasa a la cual los llamados de alarma son dados pueden también revelar la ‘urgencia’ o nivel de riesgo percibido por el emisor. En la Ratona Común, Troglodytes musculus, los individuos reproductivos dan llamadas de alarma repetitivas cuando una amenaza es detectada cerca del nido. Para entender la información que puede codificar el número de llamadas y su tasa de emisión dadas por la Ratona, testeamos la respuesta de individuos a señales experimentalmente manipuladas. Reprodujimos a campo llamados de alarma que difirieron en su tasa de repetición y duración (número de notas) y registramos la probabilidad de respuesta, la latencia de respuesta, la distancia mínima de aproximación al parlante, y el tiempo que permanecieron cerca del parlante de conespecíficos. Las Ratonas responden más frecuentemente y rápidamente a llamados de alarma repetidos a altas tasas. Además, los individuos permanecieron más tiempo cerca del parlante cuando reprodujimos llamados a tasas altas de repetición y también a secuencias más largas de llamados. Sin embargo, los individuos se aproximaron más al parlante cuando reprodujimos secuencias largas de llamados a tasas bajas. Proponemos que la repetición de las llamadas puede revelar la persistencia del estímulo, mientras que la tasa de repetición de los llamados podría codificar información acerca del nivel de riesgo detectado. Estos hallazgos contribuyen a entender la información que los receptores extraen de las señales de alarmas de coespecíficos.

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Referencias

Baker MC, Becker AM (2002) Mobbing calls of black-capped chickadees: effects of urgency on call production. The Wilson Bulletin 114(4):510-516. https://doi.org/10.1676/0043-5643(2002)114[0510:MCOBCC]2.0.CO;2
Benjamini Y, Hochberg Y (1995) Controlling the false discovery rate: a practical and powerful approach to multiple testing. Journal of the Royal Statistical Society: Series B 57(1):289-300. https://doi.org/10.1111/j.2517-6161.1995.tb02031.x
Bivand R (2022) R packages for analyzing spatial data: A comparative case study with areal data. Geographical Analysis 54(3):488-518. https://doi.org/10.1111/gean.12319
Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, Skaug HJ, Maechler M, Bolker B (2017) glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modelling. The R Journal 9(2):378-400. https://doi.org/10.32614/RJ-2017-066
Caro T (2005) Antipredator Defenses in Birds and Mammals.University of Chicago Pres, Chicago
Carro ME, Fernández GJ (2021) Alarm calls of house wrens (Troglodytes aedon bonariae) elicit responses of conspecific and heterospecific species. Behaviour 158(10):829-847. https://doi.org/10.1163/1568539X-bja10097
Common LK, Yelimlieş A, Colombelli-Négrel D, Austin VI, Kleindorfer S (2025) The ‘seet’ danger call: an active nest warning in superb fairy-wrens. Royal Society Open Science 12(11):251100. https://doi.org/10.1098/rsos.251100
Corral MG, Carro ME, Fernández GJ (2012) Alarm calls of nesting Southern house wrens (Troglodytes musculus). Ornitología Neotropical 23(2):251-260
Curio E (1978) The adaptive significance of avian mobbing: I. Teleonomic hypotheses and predictions. Zeitschrift für Tierpsychologie 48(2):175-183. https://doi.org/10.1111/j.1439-0310.1978.tb00254.x
Fallow PM, Magrath RD (2010) Eavesdropping on other species: mutual interspecific understanding of urgency information in avian alarm calls. Animal Behaviour 79(2):411-417. https://doi.org/10.1016/j.anbehav.2009.11.018
Fasanella M, Fernández GJ (2009) Alarm calls of the Southern House Wren Troglodytes musculus: variation with nesting stage and predator model. Journal of Ornithology 150(4):853-863. https://doi.org/10.1007/s10336-009-0406-2
Fernández GJ, Corral MG, Carro ME (2012) Variation in the alarm calls of Southern House Wrens (Troglodytes musculus). Emu 112(1):71-75. https://doi.org/10.1071/MU11033
Fernández GJ, Carro ME (2022) Alarm calls of southern house wrens, Troglodytes aedon bonariae, convey information about the level of risk. Ethology 128(4):293-302. https://doi.org/10.1111/eth.13255
Fernández GJ, Dutour M, Carro ME (2023) Information transfer during mobbing: call rate is more important than the number of callers in a southern temperate passerine. Behavioral Ecology and Sociobiology 77(7):78. https://doi.org/10.1007/s00265-023-03357-z
Fernández GJ, Carro ME, Johnson LS (2024) Southern House Wren (Troglodytes musculus), version 1.0. In: Juárez R, Keeney BK, Billerman S M (eds.) Birds of the World. (Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.houwre4.01
Fishbein AR, Lawson SL, Dooling RJ, Ball GF (2019) How canaries listen to their song: Species-specific shape of auditory perception. Journal of the Acoustical Society of America 145(1):562-574. https://doi.org/10.1121/1.5087692
Fox J, Weisberg S (2019) An R Companion to Applied Regression. Sage, Thousand Oaks, California
Guilford T, Dawkins MS (1991) Receiver psychology and the evolution of animal signals. Animal Behaviour 42(1):1-14. https://doi.org/10.1016/S0003-3472(05)80600-1
Harris MA, Murie JO, Duncan JA (1983) Responses of Columbian ground squirrels to playback of recorded calls. Zeitschrift für Tierpsychologie 63(4):318-330. https://doi.org/10.1111/j.1439-0310.1983.tb00747.x
Hartig F (2020) DHARMa: residual diagnostics for hierarchical (multi-level/mixed) regression models. R package ver. 0.3.3.0. [URL: http://florianhartig.github.io/DHARMa/]
Hernán MA, Robins JM (2020) Causal inference: What if. Chapman & Hall/CRC, Boca Raton
Johnstone RA (1997) The evolution of animal signals. In: Krebs JR, Davies NB (eds.) Behavioural Ecology: An Evolutionary Approach. Pp. 155-178. Blackwell Publishing, Oxford, UK
Kalb N, Anger F, Randler C (2019) Subtle variations in mobbing calls are predator-specific in great tits (Parus major). Scientific Reports 9(1):6572. https://doi.org/10.1038/s41598-019-43087-9
Klump GM, Shalter MD (1984) Acoustic behaviour of birds and mammals in the predator context; I. Factors affecting the structure of alarm signals. II. The functional significance and evolution of alarm signals. Zeitschrift für Tierpsychologie 66(3):189-226. https://doi.org/10.1111/j.1439-0310.1984.tb01365.x
Le Roux A, Jackson T, Cherry M (2001) The effect of changing call duration and calling bouts on vigilance in Brants’ whistling rat, Parotomys brantsii. Behaviour 138(10):1287-1302. https://doi.org/10.1163/15685390152822229
Leavesley AJ, Magrath RD (2005) Communicating about danger: urgency alarm calling in a bird. Animal Behaviour 70(2):365-373. https://doi.org/10.1016/j.anbehav.2004.10.017
Leger DW, Owings DH, Boal LM (1979) Contextual information and differential responses to alarm whistles in California ground squirrels. Zeitschrift für Tierpsychologie 49(2):142-155. https://doi.org/10.1111/j.1439-0310.1979.tb00283.x
Lenth R (2020) emmeans: Estimated marginal means, aka least-squares means. R package version 1.5.1. [URL: https://CRAN.R-project.org/package=emmeans]
Loughry WJ, McDonough CM (1988) Calling and vigilance in California ground squirrels: a test of the tonic communication hypothesis. Animal Behaviour 36(5):1533-1540. https://doi.org/10.1016/S0003-3472(88)80224-0
Lüdecke D, Ben-Shachar MS, Patil I, Waggoner P, Makowski D (2021) performance: An R package for assessment, comparison and testing of statistical models. Journal of Open Source Software 6(60):3139. https://doi.org/10.21105/joss.03139
Manser MB (2001) The acoustic structure of suricates’ alarm calls varies with predator type and the level of response urgency. Proceedings of the Royal Society, London 268(1483):2315-2324. https://doi.org/10.1098/rspb.2001.1773
Marler P (1955) Characteristics of some animal calls. Nature 176(4470):6-8. https://doi.org/10.1038/176006a0
Marler P (1957) Specific distinctiveness in the communication signals of birds. Behaviour 11(1):13-39. https://doi.org/10.1163/156853956X00066
McLachlan JR, Magrath RD (2020) Speedy revelations: how alarm calls can convey rapid, reliable information about urgent danger. Proceedings of the Royal Society of London B. 287(1921):20192772. https://doi.org/10.1098/rspb.2019.2772
Morton ES (1977) On the occurrence and significance of motivation-structural rules in some bird and mammal sounds. The American Naturalist 111(981):855-869. https://doi.org/10.1086/283219
Owings H., Virginia RA (1978) Alarm calls of California ground squirrels (Spermophilus beecheyi). Zeitschrift für Tierpsychologie 46(1):58-70. https://doi.org/10.1111/j.1439-0310.1978.tb01438.x
R Core Team (2020) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. [URL: https://www.R-project.org/]
Rowe C (2013) Receiver psychology: a receiver’s perspective. Animal Behaviour 85(3):517-523. https://doi.org/10.1016/j.anbehav.2013.01.004
Schleidt WM (1973) Tonic communication: continual effects of discrete signs in animal communication systems. Journal of Theoretical Biology 42(2):359-386. https://doi.org/10.1016/0022-5193(73)90095-7
Schlenker P, Chemla E, Schel AM, Fuller J, Gautier JP, Kuhn J, Veselinović D, Arnold K, Cäsar C, Keenan S, et al. (2016) Formal monkey linguistics. Theoretical Linguistics 42(1-2):1-90. https://doi.org/10.1515/tl-2016-0001
Shah SS, Greig EI, MacLean SA, Bonter DN (2015) Risk-based alarm calling in a nonpasserine bird. Animal Behaviour 106:129-136. https://doi.org/10.1016/j.anbehav.2015.05.011
Sloan JL, Hare JF (2004) Monotony and the information content of Richardson’s ground squirrel (Spermophilus richardsonii) repeated calls: Tonic communication or signal certainty? Ethology 110(2):147-156. https://doi.org/10.1111/j.1439-0310.2003.00955.x
Sommer C, Todt D, Ostreiher R, Mundry R (2012) Urgency-related alarm calling in Arabian babblers, Turdoides squamiceps: predator distance matters in the use of alarm call types. Behaviour 149(7):755-773. https://doi.org/10.1163/1568539X-00003003
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
Suzuki TN (2011) Parental alarm calls warn nestlings about different predatory threats. Current Biology 21(1):R15-R16. https://doi.org/10.1016/j.cub.2010.11.027
Suzuki TN (2016) Semantic communication in birds: evidence from field research over the past two decades. Ecological Research 31(3):307-319. https://doi.org/10.1007/s11284-016-1339-x
Templeton CN, Greene E, Davis K (2005) Allometry of alarm calls: black-capped chickadees encode information about predator size. Science 308(5730):1934-1937. https://doi.org/10.1126/science.1108841
Therneau T (2015) Mixed effects Cox models. [URL: https://cran.r-project.org/web/packages/coxme/vignettes/coxme.pdf]
Warkentin KJ, Keeley AT, Hare JF (2001) Repetitive calls of juvenile Richardson’s ground squirrels (Spermophilus richardsonii) communicate response urgency. Canadian Journal of Zoology 79(4):569-573. https://doi.org/10.1139/z01-017
Wheatcroft D (2015) Repetition rate of calls used in multiple contexts communicates presence of predators to nestlings and adult birds. Animal Behaviour 103:35-44. https://doi.org/10.1016/j.anbehav.2015.02.009
Wiley RH (1994) Errors, exaggeration, and deception in animal communication. In Real LA (ed.) Behavioral Mechanisms in Evolutionary Ecology. Pp. 157-189. University of Chicago Press, Chicago
Wiley RH (2006) Signal detection and animal communication. Advances in the Study of Behaviour 36:217-247. https://doi.org/10.1016/s0065-3454(06)36005-6
Yorzinski J L, Vehrencamp SL (2009) The effect of predator type and danger level on the mob calls of the American Crow. Condor 111(1):159-168. https://doi.org/10.1525/cond.2009.080057

Cómo citar

Gustavo J. Fernández, Mariana E. Carro (2026). Llamadas repetitivas en la Ratona Común Sudamericana: la información codificada en el número y tasa de repetición de los llamados de alarma. El Hornero, Vol. 41 Núm. 2 (2026). https://doi.org/10.56178/eh.v41i2.1543

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[1] Baker MC, Becker AM (2002) Mobbing calls of black-capped chickadees: effects of urgency on call production. The Wilson Bulletin 114(4):510-516. https://doi.org/10.1676/0043-5643(2002)114[0510:MCOBCC]2.0.CO;2

[2] Benjamini Y, Hochberg Y (1995) Controlling the false discovery rate: a practical and powerful approach to multiple testing. Journal of the Royal Statistical Society: Series B 57(1):289-300. https://doi.org/10.1111/j.2517-6161.1995.tb02031.x

[3] Bivand R (2022) R packages for analyzing spatial data: A comparative case study with areal data. Geographical Analysis 54(3):488-518. https://doi.org/10.1111/gean.12319

[4] Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, Skaug HJ, Maechler M, Bolker B (2017) glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modelling. The R Journal 9(2):378-400. https://doi.org/10.32614/RJ-2017-066

[5] Caro T (2005) Antipredator Defenses in Birds and Mammals.University of Chicago Pres, Chicago

[6] Carro ME, Fernández GJ (2021) Alarm calls of house wrens (Troglodytes aedon bonariae) elicit responses of conspecific and heterospecific species. Behaviour 158(10):829-847. https://doi.org/10.1163/1568539X-bja10097

[7] Common LK, Yelimlieş A, Colombelli-Négrel D, Austin VI, Kleindorfer S (2025) The ‘seet’ danger call: an active nest warning in superb fairy-wrens. Royal Society Open Science 12(11):251100. https://doi.org/10.1098/rsos.251100

[8] Corral MG, Carro ME, Fernández GJ (2012) Alarm calls of nesting Southern house wrens (Troglodytes musculus). Ornitología Neotropical 23(2):251-260

[9] Curio E (1978) The adaptive significance of avian mobbing: I. Teleonomic hypotheses and predictions. Zeitschrift für Tierpsychologie 48(2):175-183. https://doi.org/10.1111/j.1439-0310.1978.tb00254.x

[10] Fallow PM, Magrath RD (2010) Eavesdropping on other species: mutual interspecific understanding of urgency information in avian alarm calls. Animal Behaviour 79(2):411-417. https://doi.org/10.1016/j.anbehav.2009.11.018

[11] Fasanella M, Fernández GJ (2009) Alarm calls of the Southern House Wren Troglodytes musculus: variation with nesting stage and predator model. Journal of Ornithology 150(4):853-863. https://doi.org/10.1007/s10336-009-0406-2

[12] Fernández GJ, Corral MG, Carro ME (2012) Variation in the alarm calls of Southern House Wrens (Troglodytes musculus). Emu 112(1):71-75. https://doi.org/10.1071/MU11033

[13] Fernández GJ, Carro ME (2022) Alarm calls of southern house wrens, Troglodytes aedon bonariae, convey information about the level of risk. Ethology 128(4):293-302. https://doi.org/10.1111/eth.13255

[14] Fernández GJ, Dutour M, Carro ME (2023) Information transfer during mobbing: call rate is more important than the number of callers in a southern temperate passerine. Behavioral Ecology and Sociobiology 77(7):78. https://doi.org/10.1007/s00265-023-03357-z

[15] Fernández GJ, Carro ME, Johnson LS (2024) Southern House Wren (Troglodytes musculus), version 1.0. In: Juárez R, Keeney BK, Billerman S M (eds.) Birds of the World. (Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.houwre4.01

[16] Fishbein AR, Lawson SL, Dooling RJ, Ball GF (2019) How canaries listen to their song: Species-specific shape of auditory perception. Journal of the Acoustical Society of America 145(1):562-574. https://doi.org/10.1121/1.5087692

[17] Fox J, Weisberg S (2019) An R Companion to Applied Regression. Sage, Thousand Oaks, California

[18] Guilford T, Dawkins MS (1991) Receiver psychology and the evolution of animal signals. Animal Behaviour 42(1):1-14. https://doi.org/10.1016/S0003-3472(05)80600-1

[19] Harris MA, Murie JO, Duncan JA (1983) Responses of Columbian ground squirrels to playback of recorded calls. Zeitschrift für Tierpsychologie 63(4):318-330. https://doi.org/10.1111/j.1439-0310.1983.tb00747.x

[20] Hartig F (2020) DHARMa: residual diagnostics for hierarchical (multi-level/mixed) regression models. R package ver. 0.3.3.0. [URL: http://florianhartig.github.io/DHARMa/]

[21] Hernán MA, Robins JM (2020) Causal inference: What if. Chapman & Hall/CRC, Boca Raton

[22] Johnstone RA (1997) The evolution of animal signals. In: Krebs JR, Davies NB (eds.) Behavioural Ecology: An Evolutionary Approach. Pp. 155-178. Blackwell Publishing, Oxford, UK

[23] Kalb N, Anger F, Randler C (2019) Subtle variations in mobbing calls are predator-specific in great tits (Parus major). Scientific Reports 9(1):6572. https://doi.org/10.1038/s41598-019-43087-9

[24] Klump GM, Shalter MD (1984) Acoustic behaviour of birds and mammals in the predator context; I. Factors affecting the structure of alarm signals. II. The functional significance and evolution of alarm signals. Zeitschrift für Tierpsychologie 66(3):189-226. https://doi.org/10.1111/j.1439-0310.1984.tb01365.x

[25] Le Roux A, Jackson T, Cherry M (2001) The effect of changing call duration and calling bouts on vigilance in Brants’ whistling rat, Parotomys brantsii. Behaviour 138(10):1287-1302. https://doi.org/10.1163/15685390152822229

[26] Leavesley AJ, Magrath RD (2005) Communicating about danger: urgency alarm calling in a bird. Animal Behaviour 70(2):365-373. https://doi.org/10.1016/j.anbehav.2004.10.017

[27] Leger DW, Owings DH, Boal LM (1979) Contextual information and differential responses to alarm whistles in California ground squirrels. Zeitschrift für Tierpsychologie 49(2):142-155. https://doi.org/10.1111/j.1439-0310.1979.tb00283.x

[28] Lenth R (2020) emmeans: Estimated marginal means, aka least-squares means. R package version 1.5.1. [URL: https://CRAN.R-project.org/package=emmeans]

[29] Loughry WJ, McDonough CM (1988) Calling and vigilance in California ground squirrels: a test of the tonic communication hypothesis. Animal Behaviour 36(5):1533-1540. https://doi.org/10.1016/S0003-3472(88)80224-0

[30] Lüdecke D, Ben-Shachar MS, Patil I, Waggoner P, Makowski D (2021) performance: An R package for assessment, comparison and testing of statistical models. Journal of Open Source Software 6(60):3139. https://doi.org/10.21105/joss.03139

[31] Manser MB (2001) The acoustic structure of suricates’ alarm calls varies with predator type and the level of response urgency. Proceedings of the Royal Society, London 268(1483):2315-2324. https://doi.org/10.1098/rspb.2001.1773

[32] Marler P (1955) Characteristics of some animal calls. Nature 176(4470):6-8. https://doi.org/10.1038/176006a0

[33] Marler P (1957) Specific distinctiveness in the communication signals of birds. Behaviour 11(1):13-39. https://doi.org/10.1163/156853956X00066

[34] McLachlan JR, Magrath RD (2020) Speedy revelations: how alarm calls can convey rapid, reliable information about urgent danger. Proceedings of the Royal Society of London B. 287(1921):20192772. https://doi.org/10.1098/rspb.2019.2772

[35] Morton ES (1977) On the occurrence and significance of motivation-structural rules in some bird and mammal sounds. The American Naturalist 111(981):855-869. https://doi.org/10.1086/283219

[36] Owings H., Virginia RA (1978) Alarm calls of California ground squirrels (Spermophilus beecheyi). Zeitschrift für Tierpsychologie 46(1):58-70. https://doi.org/10.1111/j.1439-0310.1978.tb01438.x

[37] R Core Team (2020) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. [URL: https://www.R-project.org/]

[38] Rowe C (2013) Receiver psychology: a receiver’s perspective. Animal Behaviour 85(3):517-523. https://doi.org/10.1016/j.anbehav.2013.01.004

[39] Schleidt WM (1973) Tonic communication: continual effects of discrete signs in animal communication systems. Journal of Theoretical Biology 42(2):359-386. https://doi.org/10.1016/0022-5193(73)90095-7

[40] Schlenker P, Chemla E, Schel AM, Fuller J, Gautier JP, Kuhn J, Veselinović D, Arnold K, Cäsar C, Keenan S, et al. (2016) Formal monkey linguistics. Theoretical Linguistics 42(1-2):1-90. https://doi.org/10.1515/tl-2016-0001

[41] Shah SS, Greig EI, MacLean SA, Bonter DN (2015) Risk-based alarm calling in a nonpasserine bird. Animal Behaviour 106:129-136. https://doi.org/10.1016/j.anbehav.2015.05.011

[42] Sloan JL, Hare JF (2004) Monotony and the information content of Richardson’s ground squirrel (Spermophilus richardsonii) repeated calls: Tonic communication or signal certainty? Ethology 110(2):147-156. https://doi.org/10.1111/j.1439-0310.2003.00955.x

[43] Sommer C, Todt D, Ostreiher R, Mundry R (2012) Urgency-related alarm calling in Arabian babblers, Turdoides squamiceps: predator distance matters in the use of alarm call types. Behaviour 149(7):755-773. https://doi.org/10.1163/1568539X-00003003

[44] 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

[45] Suzuki TN (2011) Parental alarm calls warn nestlings about different predatory threats. Current Biology 21(1):R15-R16. https://doi.org/10.1016/j.cub.2010.11.027

[46] Suzuki TN (2016) Semantic communication in birds: evidence from field research over the past two decades. Ecological Research 31(3):307-319. https://doi.org/10.1007/s11284-016-1339-x

[47] Templeton CN, Greene E, Davis K (2005) Allometry of alarm calls: black-capped chickadees encode information about predator size. Science 308(5730):1934-1937. https://doi.org/10.1126/science.1108841

[48] Therneau T (2015) Mixed effects Cox models. [URL: https://cran.r-project.org/web/packages/coxme/vignettes/coxme.pdf]

[49] Warkentin KJ, Keeley AT, Hare JF (2001) Repetitive calls of juvenile Richardson’s ground squirrels (Spermophilus richardsonii) communicate response urgency. Canadian Journal of Zoology 79(4):569-573. https://doi.org/10.1139/z01-017

[50] Wheatcroft D (2015) Repetition rate of calls used in multiple contexts communicates presence of predators to nestlings and adult birds. Animal Behaviour 103:35-44. https://doi.org/10.1016/j.anbehav.2015.02.009

[51] Wiley RH (1994) Errors, exaggeration, and deception in animal communication. In Real LA (ed.) Behavioral Mechanisms in Evolutionary Ecology. Pp. 157-189. University of Chicago Press, Chicago

[52] Wiley RH (2006) Signal detection and animal communication. Advances in the Study of Behaviour 36:217-247. https://doi.org/10.1016/s0065-3454(06)36005-6

[53] Yorzinski J L, Vehrencamp SL (2009) The effect of predator type and danger level on the mob calls of the American Crow. Condor 111(1):159-168. https://doi.org/10.1525/cond.2009.080057