Abstract
Among territorial birds, the number of conspecific neighbours sets the social context of communication. There have been many investigations of vocal behaviour and its important role in territory defense and mate attraction in birds; however, the effect of the density of conspecific neighbours on avian communication has received little attention. In this study, we reviewed the literature on the influence of local density on vocal communication in birds, exploring how the number of conspecific neighbours influences avian vocal behaviour, by conducting a literature review following the guidelines of the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) in two databases: Web of Science and Scopus. We also included additional articles from a more flexible search in Google Scholar. We found 19 studies of birds that included data on vocal behaviour in relation to the number of conspecific neighbours. The majority of these studies (14 of 19) showed that local density influences avian vocal behaviour. Overall, these studies revealed an effect of local density of conspecifics on song rate (expressed as song rate, syllable rate, solo song rate, or singing rate). Other vocal behaviours that were shown to vary with local density included song-type switching, song duration, peak frequency, and song variation. Most studies focused on temperate-zone species where males are the predominant singers. Our results reveal a bias towards the study of male songs in the context of local density, with very few studies on female song; this corresponds with a historical geographical focus on temperate birds. We present experimental design suggestions for future investigations, including predictions for males and females of tropical and temperate species. Finally, we argue that more research on this topic is needed, and that investigations of vocal communication will benefit from further study of the influence of density on avian vocal behaviour.
Zusammenfassung
Verhaltenskonsequenzen von Artgenossen-Nachbarn: Eine systematische Literaturübersicht über die Auswirkungen der lokalen Dichte auf die vokale Kommunikation von Vögeln. In territorialen Vögeln setzt die Anzahl konspezifischer Artgenossen in der Nachbarschaft dessen Gesangsverhalten fest. Obwohl es diesbezüglich viele Untersuchungen im Zusammenhang mit Revierverteidigung und Partnerattraktion bei Vögeln gibt, wurde der Einfluss der Dichte von konspezifischen Artgenossen auf den Vogelgesang wenig beachtet. Diese Studie untersuchte die Literatur über den Einfluss der lokalen Dichte auf das Gesangsverhalten von Vögeln und den Einfluss der Anzahl konspezifischer Arten in der Nachbarschaft auf deren Kommunikation. Die Literaturstudie wurde nach den Richtlinien des Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) in zwei Datenbanken durchgeführt: Web of Science und Scopus. Wir haben auch zusätzliche Artikel aus einer flexibleren Suche in Google Scholar eingefügt. Wir fanden neunzehn Studien über das Gesangsverhalten von Vögeln im Zusammenhang mit der Anzahl von lokalen konspezifischen Artgenossen. Die meisten Studien (14 von 19) zeigten, dass die lokale Dichte die Gesangsrate (ausgedrückt als Gesangsrate, Silbenrate, Sologesangsrate oder Singrate) von Vögeln beeinflusst. Andere Gesangsverhalten, die mit der lokalen Dichte variierten, umfassten den Wechsel von Gesangstypen, die Gesangsdauer, die Höchstfrequenz und die Gesangsvariation. Die meisten Studien konzentrierten sich auf Arten der gemäßigten Zone, wo überwiegend Männchen singen. Daraus resultierend ergaben unsere Ergebnisse eine Bevorzugung der Untersuchung des Gesangsverhaltens männlicher Vögel im Kontext der lokalen Dichte durch den geografischen Fokus wissenschaftlicher Artikel auf Vögel der gemäßigten Zonen. Mit dieser Studie stellen wir zukünftige Untersuchungsvorschläge, einschließlich Vorhersagen für Männchen und Weibchen tropischer und gemäßigter Vogelarten. Weitere Forschung zu diesem Thema ist notwendig, da Studien zu dem Gesangsverhalten von Vögeln von weiteren Untersuchungen zu dem Einfluss der lokalen Dichte auf den Vogelgesang profitieren würden.
Similar content being viewed by others
Data availability
The data are included in the manuscript.
References
Alatalo RV, Glynn C, Lundberg A (1990) Singing rate and female attraction in the Pied Flycatcher: an experiment. Anim Behav 39:601–603. https://doi.org/10.1016/S0003-3472(05)80430-0
Austin VI, Dalziell AH, Langmore NE, Welbergen JA (2021) Avian vocalisations: the female perspective. Biol Rev 96:1484–1503. https://doi.org/10.1111/brv.12713
Ballentine B, Hyman J, Nowicki S (2004) Vocal performance influences female response to male bird song: an experimental test. Behav Ecol 15:163–168. https://doi.org/10.1093/beheco/arg090
Barrero A, Gómez-Catasús J, Pérez-Granados C, Bustillo-de la Rosa D, Traba J (2023) Conspecific density and habitat quality drive the defence and vocal behaviour of a territorial passerine. Ibis. https://doi.org/10.1111/ibi.13295
Beecher MD, Campbell SE, Burt JM, Hill CE, Nordby JC (2000) Song-type matching between neighbouring song sparrows. Anim Behav 59:21–27. https://doi.org/10.1006/anbe.1999.1276
Blumstein DT, Mcclain DR, De Jesus C, Alarcón-Nieto G (2012) Breeding bird density does not drive vocal individuality. Curr Zool 58:765–772. https://doi.org/10.1093/czoolo/58.5.765
Burt JM, Vehrencamp SL (2005) Dawn chorus as an interactive communication network. In: McGregor P (ed) Animal communication networks. Cambridge University Press, Cambridge, pp 320–343. https://doi.org/10.1017/CBO9780511610363.019
Cain KE, Langmore NE (2016) Female song and aggression show contrasting relationships to reproductive success when habitat quality differs. Behav Ecol Sociobio 70:1867–1877. https://doi.org/10.1007/s00265-016-2192-1
Catchpole CK, Slater PJ (2003) Bird song: biological themes and variations. Cambridge University Press
Christman SP (1984) Plot mapping: estimating densities of breeding bird territories by combining spot mapping and transect techniques. Condor 86:237–241
Connor EF, Courtney AC, Yoder JM (2000) Individuals–area relationships: the relationship between animal population density and area. Ecology 81:734–748
Cooney R, Cockburn A (1995) Territorial defence is the major function of female song in the superb fairy-wren, Malurus cyaneus. Anim Behav 49:1635–1647. https://doi.org/10.1016/0003-3472(95)90086-1
Dargis L, Benedict L, Najar NA (2021) Female bird song rates do not covary with population density in a North American species. Ethology 127:1042–1052. https://doi.org/10.1111/eth.13227
Dawson DK, Efford MG (2009) Bird population density estimated from acoustic signals. J Appl Ecol 46:1201–1209. https://doi.org/10.1111/j.1365-2664.2009.01731.x
Demko AD, Mennill DJ (2018) Male and female signaling behavior varies seasonally during territorial interactions in a tropical songbird. Behav Ecol Sociobiol 72:1–13. https://doi.org/10.1007/s00265-018-2495-5
Falls JB (1981) Mapping territories with playback: an accurate census method for songbirds. Stud Avian Biol 6:86–91
Foote JR, Fitzsimmons LP, Mennill DJ, Ratcliffe LM (2008) Male chickadees match neighbors interactively at dawn: support for the social dynamics hypothesis. Behav Ecol 19:1192–1199. https://doi.org/10.1093/beheco/arn087
Foote JR, Ratcliffe LM, Mennill DJ, Fitzsimmons LP (2010) Black-capped chickadee dawn choruses are interactive communication networks. Behaviour 147:1219–1248
Franco P, Slabbekoorn H (2009) Repertoire size and composition in great tits: a flexibility test using playbacks. Anim Behav 77:261–269. https://doi.org/10.1016/j.anbehav.2008.09.023
Galeotti P, Saino N, Sacchi R, Møller AP (1997) Song correlates with social context, testosterone and body condition in male barn swallows. Anim Behav 53:687–700. https://doi.org/10.1006/anbe.1996.0304
Gaston KJ, Matter SF (2002) Individuals-area relationships: comment. Ecology 83:288–293. https://doi.org/10.2307/2680139
Goretskaia MI (2004) Song structure and singing behaviour of willow warbler Phylloscopus trochilus acredula in populations of low and high density. Bioacoustics 14:183–195. https://doi.org/10.1080/09524622.2004.9753524
Grabarczyk EE, Vonhof MJ, Gill SA (2020) Social context and noise affect within and between male song adjustments in a common passerine. Behav Ecol 31:1150–1158. https://doi.org/10.1093/beheco/araa066
Halupka K (2014) Spreading information in a network of interacting neighbours. PLoS ONE 9:e102801. https://doi.org/10.1371/journal.pone.0102801
Halupka K (2017) Information flow in a network of dispersed signalers-receivers. Phys A Stat Mech 486:610–617. https://doi.org/10.1016/j.physa.2017.05.068
Hamao S, Watanabe M, Mori Y (2011) Urban noise and male density affect songs in the great tit Parus major. Ethol Ecol Evol 23:111–119. https://doi.org/10.1080/03949370.2011.554881
Hiebert SM, Stoddard PK, Arcese P (1989) Repertoire size, territory acquisition and reproductive success in the song sparrow. Anim Behav 37:266–273. https://doi.org/10.1016/0003-3472(89)90115-2
Janicke T, Hahn S, Ritz MS, Peter HU (2008) Vocal performance reflects individual quality in a nonpasserine. Anim Behav 75:91–98
Jetz W, Thomas GH, Joy JB, Hartmann K, Mooers AO (2012) The global diversity of birds in space and time. Nature 491:444. https://doi.org/10.1016/j.anbehav.2007.04.007
Kroodsma DE (1999) Making ecological sense of song development by songbirds. In: Hauser MD, Konishi M (eds) The Design of Animal Communication. MIT Press. https://doi.org/10.7551/mitpress/2359.003.0014
Langmore NE (1998) Functions of duet and solo songs of female birds. Trends Ecol Evol 13:136–140. https://doi.org/10.1016/S0169-5347(97)01241-X
Langmore NE (2000) Why female birds sing. In: Espmark Y, Amundsen T, Rosenqvist G (eds) Animal signals: signalling and signal design in animal communication. Tapir Academic Press, Trondheim, pp 317–327
Liu WC (2004) The effect of neighbours and females on dawn and daytime singing behaviours by male chipping sparrows. Anim Behav 68:39–44. https://doi.org/10.1016/j.anbehav.2003.06.022
Logue DM (2005) Cooperative defence in duet singing birds. Cogn Brain, Behav. 10:497–510
Mayor SJ, Schaefer JA (2005) The many faces of population density. Oecologia 145:275–280. https://doi.org/10.1007/s00442-005-0114-4
McGregor PK (2005) Animal communication networks. Cambridge University Press. https://doi.org/10.1017/CBO9780511610363
Mennill DJ (2006) Aggressive responses of male and female rufous-and-white wrens to stereo duet playback. Anim Behav 71:219–226. https://doi.org/10.1016/j.anbehav.2005.05.006
Mennill DJ, Ratcliffe LM (2004) Do male black-capped chickadees eavesdrop on song contests? A multi-speaker playback experiment. Behaviour 141:125–139
Mennill DJ, Vehrencamp SL (2008) Context-dependent functions of avian duets revealed by microphone-array recordings and multi-speaker playback. Curr Biol 18:1314–1319. https://doi.org/10.1016/j.cub.2008.07.073
Mennill DJ, Ratcliffe LM, Boag PT (2002) Female eavesdropping on male song contests in songbirds. Science 296:873–873. https://doi.org/10.1126/science.296.5569.873
Mennill DJ, Burt JM, Fristrup KM, Vehrencamp SL (2006) Accuracy of an acoustic location system for monitoring the position of duetting songbirds in tropical forest. J Acoust Soc Am 119:2832–2839. https://doi.org/10.1121/1.2184988
Moher D, Liberati A, Tetzlaff J, Altman DG, Altman D, Antes G, Atkins D, Barbour V, Barrowman N, Berlin JA, Clark J (2009) Preferred reporting items for systematic reviews and meta-analyses: the PRISMA statement (Chinese edition). Chin J Integr Med 7:889–896. https://doi.org/10.1371/journal.pmed.1000097
Moiron M, González-Lagos C, Slabbekoorn H, Sol D (2015) Singing in the city: high song frequencies are no guarantee for urban success in birds. Behav Ecol 26:843–850. https://doi.org/10.1093/beheco/arv026
Naguib M (2005) Singing interactions in songbirds: Implications for social relations. In: McGregor P (ed) Animal communication networks. Cambridge University Press, Cambridge, p 300
Naguib M, Mennill DJ (2010) The signal value of birdsong: empirical evidence suggests song overlapping is a signal. Anim Behav 80:E11–E15. https://doi.org/10.1017/CBO9780511610363.018
Narango DL, Rodewald AD (2016) Urban-associated drivers of song variation along a rural–urban gradient. Behav Ecol 27:608–616. https://doi.org/10.1093/beheco/arv197
Nolan PM, Hill GE (2004) Female choice for song characteristics in the house finch. Anim Behav 67:403–410. https://doi.org/10.1016/j.anbehav.2003.03.018
Nowicki S, Searcy WA (2005) Song and mate choice in birds: how the development of behavior helps us understand function. Auk 122:1–14. https://doi.org/10.1093/auk/122.1.1
Nowicki S, Searcy W, Hughes M (1998) The territory defense function of song in song sparrows: a test with the speaker occupation design. Behaviour 135:615–628. https://doi.org/10.1163/156853998792897888
Odom KJ, Benedict L (2018) A call to document female bird songs: applications for diverse fields. Auk Ornithol Adv 135:314–325. https://doi.org/10.1642/AUK-17-183.1
Odom KJ, Hall ML, Riebel K, Omland KE, Langmore NE (2014) Female song is widespread and ancestral in songbirds. Nat Commun 5:1–6. https://doi.org/10.1038/ncomms4379
Olinkiewicz A, Osiejuk TS (2003) Effect of time of season and neighbours on singing activity in the Corn Bunting Miliaria calandra. Acta Ornithol 38:117–122. https://doi.org/10.3161/068.038.0208
Opaev AS, Shishkina EM (2021) Song amplitude and population density in two sympatric warblers, Phylloscopus schwarzi and P. fuscatus. Bioacoustics 30:272—283. https://doi.org/10.1080/09524622.2020.1720816
Osmun AE, Mennill DJ (2011) Acoustic monitoring reveals congruent patterns of territorial singing behaviour in male and female tropical wrens. Ethology 117:385–394. https://doi.org/10.1111/j.1439-0310.2011.01887.x
Owen KC, Mennill DJ (2021) Singing in a fragmented landscape: wrens in a tropical dry forest show sex differences in the effects of neighbours, time of day, and time of year. J Ornithol 162:881–893. https://doi.org/10.1007/s10336-021-01882-0
Peake TM, Terry AMR, McGregor PK, Dabelsteen T (2002) Do great tits assess rivals by combining direct experience with information gathered by eavesdropping? Proc R Soc B Biol Sci 269:1925–1929. https://doi.org/10.1098/rspb.2002.2112
Pérez-Granados C, Traba J (2021) Estimating bird density using passive acoustic monitoring: a review of methods and suggestions for further research. Ibis 163:765–783. https://doi.org/10.1111/ibi.12944
Podos J, Lahti DC, Moseley DL (2009) Vocal performance and sensorimotor learning in songbirds. Adv Study Behav 40:159–195. https://doi.org/10.1016/S0065-3454(09)40005-6
Reid JM, Arcese P, Cassidy AL, Hiebert SM, Smith JN, Stoddard PK, Marr AB, Keller LF (2004) Song repertoire size predicts initial mating success in male song sparrows, Melospiza melodia. Anim Behav 68:1055–1063. https://doi.org/10.1016/j.anbehav.2004.07.003
Riebel K (2009) Song and female mate choice in zebra finches: a review. Adv Study Behav 40:197–238. https://doi.org/10.1016/S0065-3454(09)40006-8
Riebel K, Odom KJ, Langmore NE, Hall ML (2019) New insights from female bird song: towards an integrated approach to studying male and female communication roles. Biol Lett 15:20190059. https://doi.org/10.1098/rsbl.2019.0059
Ripmeester EA, Kok JS, van Rijssel JC, Slabbekoorn H (2010) Habitat-related birdsong divergence: a multi-level study on the influence of territory density and ambient noise in European blackbirds. Behav Ecol Sociobiol 64:409–418. https://doi.org/10.1007/s00265-009-0857-8
Searcy WA (1992) Song repertoire and mate choice in birds. Am Zool 32:71–80. https://doi.org/10.1093/icb/32.1.71
Searcy WA, Andersson M (1986) Sexual selection and the evolution of song. Annu Rev Ecol Evol Syst 17:507–533. https://doi.org/10.1146/annurev.es.17.110186.002451
Searcy WA, Beecher MD (2009) Song as an aggressive signal in songbirds. Anim Behav 78:1281–1292. https://doi.org/10.1016/j.anbehav.2009.08.011
Searcy WA, Akçay C, Nowicki S, Beecher MD (2014) Aggressive signaling in song sparrows and other songbirds. Adv Study Behav 46:89–125. https://doi.org/10.1016/B978-0-12-800286-5.00003-1
Sexton K, Murphy M, Redmond L, Dolan AC (2007) Dawn song of eastern kingbirds: intrapopulation variability and sociobiological correlates. Behaviour 144:1273–1295. https://doi.org/10.1163/156853907781890922
Sharma SPS, Dobney SL, Norris DR, Doucet SM, Newman AEM, Burant JB, Moran IG, Mueller SD, Spina HA, Woodworth B, Mennill DJ (2024). Intrinsic and extrinsic factors influence territory size and shape in Savannah Sparrows: effects of age, breeding strategy, population density, and number of neighbours. Ornithol in press
Shutler D, Weatherhead PJ (1991) Basal song rate variation in male red-winged blackbirds: sound and fury signifying nothing? Behav Ecol 2:123–132. https://doi.org/10.1093/beheco/2.2.123
Sillett TS, Rodenhouse NL, Holmes RT (2004) Experimentally reducing neighbor density affects reproduction and behavior of a migratory songbird. Ecology 85:2467–2477. https://doi.org/10.1890/03-0272
Slater PJ, Mann NI (2004) Why do the females of many bird species sing in the tropics? J Avian Biol 35:289–294. https://doi.org/10.1111/j.0908-8857.2004.03392.x
Snijders L, Naguib M (2017) Communication in animal social networks: a missing link? In: Naguib M, Podos J, Simmons LW, Barrett L, Healy SD, Zuk M (eds) Advances in the study of behavior. Elsevier Academic Press, pp 297–359. https://doi.org/10.1016/bs.asb.2017.02.004
Staicer CA, Spector DA, Horn AC (1996) The dawn chorus and other diel patterns in acoustic signaling. In: Kroodsma DE, Miller EH (eds) Ecology and evolution of acoustic communication in birds. Cornell University Press, Ithaca, pp 426–453. https://doi.org/10.7591/9781501736957-033
Stehelin TE, Ross M, Lein M (2014) Social stimulation of dawn singing in Dusky Flycatchers: a serendipitous experiment. J Field Ornithol 85:63–74. https://doi.org/10.1111/jofo.12050
Stuart CJ, Grabarczyk EE, Vonhof MJ, Gill SA (2019) Social factors, not anthropogenic noise or artificial light, influence onset of dawn singing in a common songbird. Auk 136:ukz045. https://doi.org/10.1093/auk/ukz045
Stutchbury BJ, Morton ES (2001) Behavioral ecology of tropical birds. Academic Press, San Diego
Stutchbury BJ, Morton ES (2022) Behavioral ecology of tropical birds. Academic Press, San Diego
Tobias JA, Gamarra-Toledo V, García-Olaechea D, Pulgarin PC, Seddon N (2011) Year-round resource defence and the evolution of male and female song in sub-oscine birds: social armaments are mutual ornaments. J Evol Biol 24:2118–2138. https://doi.org/10.1111/j.1420-9101.2011.02345.x
Tobias JA, Sheard C, Seddon N, Meade A, Cotton AJ, Nakagawa S (2016) Territoriality, social bonds, and the evolution of communal signaling in birds. Front Ecol Evol 4:74. https://doi.org/10.3389/fevo.2016.00074
Todt D, Naguib M (2000) Vocal interactions in birds: the use of song as a model in communication. Advances in the study of behavior, vol 29. Academic Press, pp 247–296. https://doi.org/10.1016/S0065-3454(08)60107-2
Valderrama SV, Molles LE, Waas JR (2013) Effects of population size on singing behavior of a rare duetting songbird. Conserv Biol 27:210–218. https://doi.org/10.1111/j.1523-1739.2012.01917.x
Vehrencamp SL (2001) Is song–type matching a conventional signal of aggressive intentions? Proc R Soc B Biol Sci 268:1637–1642. https://doi.org/10.1098/rspb.2001.1714
Voigt C, Leitner S, Gahr M, Ter Maat A (2021) Seasonal and diurnal variation of vocal behaviour in duetting white-browed sparrow weavers. J Ornithol 162:1163–1172. https://doi.org/10.1007/s10336-021-01905-w
Wilkins MR, Odom KJ, Benedict L, Safran RJ (2020) Analysis of female song provides insight into the evolution of sex differences in a widely studied songbird. Anim Behav 168:69–82. https://doi.org/10.1016/j.anbehav.2020.07.018
Xia C, Wei C, Lloyd H, Liu J, Wu Q, Zhang Y (2014) Dawn singing intensity of the Male Brownish-Flanked Bush Warbler: effects of territorial insertions and number of neighbors. Ethology 120:324–330. https://doi.org/10.1111/eth.12205
Acknowledgements
The authors thank Mitacs Canada, and supporting partners Birds Canada and Wildlife Acoustics, for a grant supporting NVS. The authors thank the Guanacaste Dry Forest Conservation Fund for support to NVS. The authors thank NSERC for Discovery Grant support to DJM. The authors thank Kian Lanzenstiel for translating the abstract into German. The authors thank J. Aubin, S. Dobney, and D. Zapata for comments on an early version of this manuscript, and two anonymous reviewers who provided thoughtful feedback that improved the manuscript.
Author information
Authors and Affiliations
Contributions
NVS conceived of the idea, design the study, collected the data, wrote the first version of the manuscript, and edited subsequent versions of the manuscript. DJM conceived of the idea, and wrote and edited multiple versions of the manuscript.
Corresponding author
Ethics declarations
Conflict of interest
We declare no conflict of interest.
Additional information
Communicated by T. S. Osiejuk.
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Rights and permissions
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.
About this article
Cite this article
Sánchez, N.V., Mennill, D.J. Behavioural consequences of conspecific neighbours: a systematic literature review of the effects of local density on avian vocal communication. J Ornithol (2024). https://doi.org/10.1007/s10336-024-02188-7
Received:
Revised:
Accepted:
Published:
DOI: https://doi.org/10.1007/s10336-024-02188-7