Abstract
Low-amplitude signals function in private exchanges of information between signalers and nearby receivers. The eavesdropping avoidance hypothesis proposes that selection favors quiet threat signals in order to avoid the costs of eavesdroppers. If true, then selection should favor other acoustic traits in addition to low amplitude that lead to quiet signals transmitting less effectively through the environment compared to broadcast signals. The “warbled” soft songs of male song sparrows differ from “crystallized” soft songs and from broadcast songs in a number of acoustic traits, suggesting that these songs may transmit less effectively. We tested this prediction in a field experiment by playing broadcast songs, crystallized soft songs, and warbled soft songs through a loudspeaker at the same amplitude and recording the propagated songs at five distances, at two heights, and in two different habitat types. Counter to our prediction, we found no evidence that either form of soft song transmits differently than broadcast song when all were played loudly. If anything, soft songs transmitted more effectively when all songs were played quietly. Our results do not support one prediction made by the eavesdropping avoidance hypothesis, although the possibility remains that reduced amplitude alone is sufficient to reduce eavesdropping. The question of why warbled soft song differs in acoustic structure remains unresolved.
Significance statement
Quiet aggressive signals are a puzzle because their meek form seems counter to their purpose—to threaten and intimidate rivals. One explanation for quiet signals is that reducing the signal’s transmission range reduces the costs imposed by eavesdroppers, which predicts that quiet signals will transmit through the environment poorly relative to louder, broadcast signals. We tested this prediction by playing song sparrow “soft songs” and “broadcast songs” at the same amplitudes and measured their transmission properties at different distances and heights, and in two habitats. We found that soft and broadcast songs did not transmit differently when played loudly, and soft songs transmitted slightly better when played softly. Our results do not support the idea that quiet signals are explained by eavesdropping avoidance, at least not in the song sparrow.
Similar content being viewed by others
References
Akaike H (1974) New look at statistical model identification. IEEE Trans Autom Control 19:716–723
Akçay Ç, Anderson RC, Nowicki S, Beecher MD, Searcy WA (2015) Quiet threats: soft song as an aggressive signal in birds. Anim Behav 105:267–274
Akçay Ç, Beecher MD (2012) Signaling while fighting: further comments on soft song. Anim Behav 83:e1–e3
Akçay Ç, Clay A, Campbell SE, Beecher MD (2016) The sparrow and the hawk: aggressive signaling under risk of predation. Behav Ecol 27:601–607
Akçay Ç, Tom ME, Holmes D, Campbell SE, Beecher MD (2011) Sing softly and carry a big stick: signals of aggressive intent in the song sparrow. Anim Behav 82:377–382
Anderson RC, Searcy WA, Peters S, Nowicki S (2008) Soft song in song sparrows: acoustic structure and implications for signal function. Ethology 114:662–676
Ballentine B, Searcy WA, Nowicki S (2008) Reliable aggressive signalling in swamp sparrows. Anim Behav 75:693–703
Balsby TJS, Dabelsteen T, Pedersen SB (2003) Degradation of whitethroat vocalisations: implications for song flight and communication network activities. Behaviour 140:695–719
Bolt LM (2014) Male-specific use of the purr in the ring-tailed lemur (Lemur catta). Folia Primatol 85:201–214
Boogert NJ, Anderson RC, Peters S, Searcy WA, Nowicki S (2011) Song repertoire size in male song sparrows correlates with detour reaching, but not with other cognitive measures. Anim Behav 81:1209–1216
Bourne GR, Collins AC, Holder AM, McCarthy CL (2001) Vocal communication and reproductive behavior of the frog Colostethus beebei in Guyana. J Herpetol 35:272–281
Clarke E, Reichard UH, Zuberbuhler K (2015) Context-specific close-range “hoo” calls in wild gibbons (Hylobates lar). BMC Evol Biol 15:56
Dabelsteen T (2005) Public, private or anonymous? Facilitating and countering eavesdropping. In: McGregor PK (ed) Animal communication networks. Cambridge University Press, Cambridge, pp 38–62
Dabelsteen T, Larsen ON, Pedersen SB (1993) Habitat-induced degradation of sound signals—quantifying the effects of communication sounds and bird location on blur ratio, excess attenuation, and signal-to-noise ratio in blackbird song. J Acoust Soc Am 93:2206–2220
Hof D, Hazlett N (2010) Low-amplitude song predicts attack in a North American wood warbler. Anim Behav 80:821–828
Holland J, Dabelsteen T, Bjørn C, Pedersen S (2001) The location of ranging cues in wren song: evidence from calibrated interactive playback experiments. Behaviour 138:189–206
Jakobsson S, Brick O, Kullberg C (1995) Escalated fighting behaviour incurs increased predation risk. Anim Behav 49:235–239
Krams I (2001) Communication in crested tits and the risk of predation. Anim Behav 61:1065–1068
Ladich F (2007) Females whisper briefly during sex: context- and sex-specific differences in sounds made by croaking gouramis. Anim Behav 73:379–387
Marten K, Marler P (1977) Sound-transmission and its significance for animal vocalization. 1. Temperate habitats. Behav Ecol Sociobiol 2:271–290
McGregor PK (1993) Signalling in territorial systems: a context for individual identification, ranging and eavesdropping. Philos Trans R Soc B 340:237–244
McGregor PK, Dabelsteen T (1996) Communication networks. In: Kroodsma D, Miller E (eds) Ecology and evolution of acoustic communication in birds. Cornell University Press, New York, pp 409–425
Mennill DJ, Ratcliffe LM, Boag PT (2002) Female eavesdropping on male song contests in songbirds. Science 296:873–873
Morton ES (1975) Ecological sources of selection on avian sounds. Am Nat 109:17–34
Mougeot F, Bretagnolle V (2000) Predation as a cost of sexual communication in nocturnal seabirds: an experimental approach using acoustic signals. Anim Behav 60:647–656
Nice M (1943) Studies in the life history of the song sparrow II. The behavior of the song sparrow and other passerines. Trans Linn Soc N Y 6:1–328
Peters S, Searcy WA, Beecher MD, Nowicki S (2000) Geographic variation in the organization of song sparrow repertoires. Auk 117:936–942
Reichard DG, Anderson RC (2015) Why signal softly? The structure, function and evolutionary significance of low-amplitude signals. Anim Behav 105:253–265
Reichard DG, Rice RJ, Schultz EM, Schrock SE (2013) Low-amplitude songs produced by male dark-eyed juncos (Junco hyemalis) differ when sung during intra- and inter-sexual interactions. Behaviour 150:1183–1202
Reichard DG, Rice RJ, Vanderbilt CC, Ketterson ED (2011) Deciphering information encoded in birdsong: male songbirds with fertile mates respond most strongly to complex, low-amplitude songs used in courtship. Am Nat 178:478–487
Reichard DG, Welklin JF (2015) On the existence and potential functions of low-amplitude vocalizations in North American birds. Auk 132:156–166
Rek P (2013) Soft calls and broadcast calls in the corncrake as adaptations to short and long range communication. Behav Process 99:121–129
Rek P, Osiejuk TS, Budka M (2011) Functionally similar acoustic signals in the corncrake (Crex crex) transmit information about different states of the sender during aggressive interactions. Horm Behav 60:706–712
Ritschard M, Riebel K, Brumm H (2010) Female zebra finches prefer high-amplitude song. Anim Behav 79:877–883
Ritschard M, van Oers K, Naguib M, Brumm H (2012) Song amplitude of rival males modulates the territorial behaviour of great tits during the fertile period of their mates. Ethology 118:197–202
Ryan MJ, Tuttle MD, Rand AS (1982) Bat predation and sexual advertisement in a neotropical anuran. Am Nat 119:136–139
Searcy WA, Anderson RC, Nowicki S (2006) Bird song as a signal of aggressive intent. Behav Ecol Sociobiol 60:234–241
Searcy WA, Nowicki S (2006) Signal interception and the use of soft song in aggressive interactions. Ethology 112:865–872
Searcy WA, Yasukawa K (2017) Eavesdropping and cue denial in avian acoustic signals. Anim Behav 124:273–282
Vargas-Castro LE, Sandoval L, Searcy WA (2017) Eavesdropping avoidance and sound propagation: the acoustic structure of soft song. Anim Behav 134:113–121
Titus RC (1998) Short-range and long-range songs: use of two acoustically distinct song classes by dark-eyed juncos. Auk 115:386–393
Xia C, Liu J, Alström P, Wu Q, Zhang Y, Janik V (2013) Is the soft song of the brownish-flanked bush warbler an aggressive signal? Ethology 119:653–661
Zollinger SA, Brumm H (2015) Why birds sing loud songs and why they sometimes don't. Animal Behaviour 105:289–295
Acknowledgments
We thank Caitlin Cantrell for help with field work. We thank Florida Atlantic University for funds to RA and the Duke Office of the Provost for funds to SN. Access to field sites was granted by the Pennsylvania State Game Commission, and logistical support in the field was provided by the Pymatuning Laboratory of Ecology of the University of Pittsburgh. We also thank two anonymous reviewers whose suggestions improved the manuscript.
Author information
Authors and Affiliations
Corresponding author
Ethics declarations
Ethical approval
Our study did not involve animal subjects, only playback and recording of audio that had been collected in a previous study.
Conflict of interest
The authors declare that they have no conflict of interest.
Additional information
Communicated by B. Voelkl
Rights and permissions
About this article
Cite this article
Niederhauser, J.M., DuBois, A.L., Searcy, W.A. et al. A test of the eavesdropping avoidance hypothesis as an explanation for the structure of low-amplitude aggressive signals in the song sparrow. Behav Ecol Sociobiol 72, 47 (2018). https://doi.org/10.1007/s00265-018-2469-7
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s00265-018-2469-7