The number of syllables per phrase in Great Tit (Parus major) song decreases in strong anthropogenic noise and at northern latitudes


  • Magne Husby Nord University
  • Tore Slagsvold



anthropogenic noise, bird communication, bird song, highway noise, latitude


Cover photo: Great Tit male. Photo: Alf Tore Mjøs.

Birds may sing to defend a territory and to attract a mate. However, despite many studies clear conclusions remain on how ecological conditions affect the song, such as physical obstacles that may reduce the sound transmission, and anthropogenic noise that may mask the signal. The social environment of the local populations may also be important, such as breeding density and sex ratio, influencing the number of competing males with which to song match, and the distances to the neighbouring males and to prospecting females. During 2016-19, we counted the number of syllables (notes) per phrase of singing male Great Tits Parus major by visiting seven countries in Europe and one in North Africa. A total of 946 songs were observed by visiting 554 territories. We also recorded study year, anthropogenic noise, calendar date, time of day, type of habitat and vegetation density, latitude, longitude and altitude. The most important explanatory variables were anthropogenic noise and the latitude of the focal site; the number of syllables per phrase decreasing both with increasing anthropogenic noise and with the latitude. The latter result was also supported when analysing sonograms of the species found on the Internet (Xeno-canto), namely fewer syllables per phrase in Norway than in Spain and Portugal. We suggest that repetition of a short phrase is fast interpreted by conspecifics in noisy environments, and that such signals are more readily detected by conspecifics over a wider area where the density of the tits is low.


Akcay C, Porsuk YK, Avsar A, Cabuk D & Bilgin CC. 2020. Song overlapping, noise, and territorial aggression in Great Tits. Behavioral Ecology 31: 807–814.

Baker MC, Bjerke TK, Lampe H & Espmark Y. 1986. Sexual response of female Great Tits to variation in size of males’ song repertoires. American Naturalist 128: 491–498.

Bergen F & Abs M. 1997. Etho-ecological study of the singing activity of the Blue Tit (Parus caeruleus), Great Tit (Parus major) and Chaffinch (Fringilla coelebs). Journal für Ornithologie 138: 451–467.

Berger-Tal O, Wong BBM, Candolin U & Barber J. 2019. What evidence exists on the effects of anthropogenic noise on acoustic communication in animals? A systematic map protocol. Environmental Evidence 8 (Suppl 1): art18.

Bergman, G. 1980. Die Veränderung der Gesangmelodie der Kohlmeise Parus major in Finnland and Schweden (The change of song pattern of the Great Tit Parus major in Finland and Sweden). Ornis Fennica 57: 97–111.

Bermúdez-Cuamatzin E, Delamore Z, Verbeek L, Kremer C & Slabbekoorn H. 2020. Variation in diurnal patterns of singing activity between urban and rural Great Tits. Frontiers in Ecology and Evolution 8: art246.

Bermúdez-Cuamatzin E, Rios-Chelen AA, Gil D & Garcia CM. 2009. Strategies of song adaptation to urban noise in the House Finch: syllable pitch plasticity or differential syllable use? Behaviour 146: 1269–1286.

Blumenrath SH & Dabelsteen T. 2004. Degradation of Great Tit (Parus major) song before and after foliation: Implications for vocal communication in a deciduous forest. Behaviour 141: 935–958.

Bolker BM, Brooks ME, Clark CJ, Geange SW, Poulsen JR, Stevens MHH & White JSS. 2009. Generalized linear mixed models: a practical guide for ecology and evolution. Trends in Ecology & Evolution 24: 127–135.

Brumm H & Naguib M. 2009. Environmental acoustics and the evolution of bird song. Pp. 1–33 in M Naguib, K Zuberbuhler, NS Clayton & VM Janik (Eds.) Advances in the Study of Behavior , Vol. 40. Elsevier Academic Press Inc, San Diego.

Brumm H & Slabbekoorn H. 2005. Acoustic communication in noise. Pp. 151–209 in Slater PJB, Snowdon CT, Brockmann HJ, Roper TJ & Naguib M (Eds.) Advances in the Study of Behavior, Vol. 35. Elsevier Academic Press Inc, San Diego.

Brumm H & Slater PJB. 2006. Ambient noise, motor fatigue, and serial redundancy in Chaffinch song. Behavioral Ecology and Sociobiology 60: 475–481.

Brumm H & Zollinger SA. 2011. The evolution of the Lombard effect: 100 years of psychoacoustic research. Behaviour 148: 1173–1198.

Bueno-Enciso J, Ferrer ES, Barrientos R & Sanz JJ. 2016. Habitat structure influences the song characteristics within a population of Great Tits Parus major. Bird Study 63: 359–368.

Bueno-Enciso J, Nunez-Escribano D & Sanz JJ. 2015. Cultural transmission and its possible effect on urban acoustic adaptation of the Great Tit Parus major. Animal Biodiversity and Conservation 38: 221–231.

Burnham KP & Anderson DR. 2002. Model selection and multimodel inference. A practical information-theoretic approach. Second Edition. Springer, New York.

Bye FN. 2006. Kjøttmeis Parus major. Pp. 356–357 in Svorkmo-Lundberg T, Bakken V, Helberg M, Mork K, Røer JE & Sæbø S (Eds.) Norsk VinterfuglAtlas. Fuglenes utbredelse, bestandsstørrelse og økologi vinterstid. Norsk Ornitologisk Forening, Trondheim.

Byers BE & Kroodsma DE. 2009. Female mate choice and songbird song repertoires. Animal Behaviour 77: 13–22.

Cardoso GC, Klingbeil BT, La Sorte FA, Lepczyk CA, Fink D & Flather CH. 2020. Exposure to noise pollution across North American passerines supports the noise filter hypothesis. Global Ecology and Biogeography 29: 1430–1434.

Cartwright LA, Taylor DR, Wilson DR & Chow-Fraser P. 2014. Urban noise affects song structure and daily patterns of song production in Red-winged Blackbirds (Agelaius phoeniceus). Urban Ecosystems 17: 561–572.

Catchpole CK & Slater PJB. 2008. Bird song: biological themes and variations. Cambridge University Press, Cambridge.

Colino-Rabanal VJ, Mendes S, Peris SJ & Pescador M. 2016. Does the song of the wren Troglodytes troglodytes change with different environmental sounds? Acta Ornithologica 51: 13–22.

Collins SA, de Kort SR, Perez-Tris J & Telleria JL. 2009. Migration strategy and divergent sexual selection on bird song. Proc R Soc B 276: 585–590.

Cramp S & Perrins CM. 1993. The birds of the Western Palearctic. Vol. 7: Flycatchers to shrikes. Oxford University Press, Oxford.

Crouch NMA & Mason-Gamer RJ. 2019. Identifying ecological drivers of interspecific variation in song complexity in songbirds (Passeriformes, Passeri). Journal of Avian Biology 50: 14.

Dabelsteen T, McGregor PK, Shepherd M, Whittaker X & Pedersen SB. 1996. Is the signal value of overlapping different from that of alternating during matched singing in Great Tits? Journal of Avian Biology 27: 189–194.

Deoniziak K & Osiejuk TS. 2016. Disentangling relations among repertoire size, song rate, signal redundancy and ambient noise level in European songbird. Ethology 122: 734–744.

Deoniziak K & Osiejuk TS. 2019. Habitat-related differences in song structure and complexity in a songbird with a large repertoire. BMC Ecology 19: art40.

Deoniziak K & Osiejuk TS. 2021. Seasonality and social factors, but not noise pollution, influence the song characteristics of two leaf warbler species. Plos One 16: e0257074.

Dormann CF, Elith J, Bacher S, Buchmann CCG, Carre G, Marquez JRG, Gruber B, Lafourcade B, Leitao PJ, Munkemuller T, McClean C, Osborne PE, Reineking B, Schroder B, Skidmore AK, Zurell D & Lautenbach S. 2013. Collinearity: a review of methods to deal with it and a simulation study evaluating their performance. Ecography 36: 27–46.

Dowling JL, Luther DA & Marra PP. 2012. Comparative effects of urban development and anthropogenic noise on bird songs. Behavioral Ecology 23: 201–209.

Emlen ST. 1972. An experimental analysis of the parameters of bird song eliciting species recognition. Behaviour 41: 130–171.

Ey E & Fischer J. 2012. The ’acoustic adaptation hypthesis’ – a review of the evidence from birds, anurans and mammals. Bioacoustics 19: 21–48.

Fernández-Juricic E, Poston R, De Collibus K, Morgan T, Bastain B, Martin C, Jones K & Treminio R. 2005. Microhabitat selection and singing behavior patterns of male House Finches (Carpodacus mexicanus) in urban parks in a heavily urbanized landscape in the western U.S. Urban Habitats 3: 49–69.

Ferrer M, de Lucas M, Janss GFE, Casado E, Munoz AR, Bechard MJ & Calabuig CP. 2012. Weak relationship between risk assessment studies and recorded mortality in wind farms. Journal of Applied Ecology 49: 38–46.

Foppen R & Reijnen R. 1994. The effects of car traffic on breeding bird populations in woodland. 2. Breeding dispersal of male Willow Warblers (Phylloscopus trochilus) in relation to the proximity of a highway. Journal of Applied Ecology 31: 95–101.

Forman RTT 2000. Estimate of the area affected ecologically by the road system in the United States. Conservation Biology 14: 31–35.

Forman RTT & Alexander LE. 1998. Roads and their major ecological effects. Annual Review of Ecology and Systematics 29: 207–231.

Francis CD, Ortega CP & Cruz A. 2009. Noise pollution changes avian communities and species interactions. Current Biology 19: 1415–1419.

Franco P & Slabbekoorn H. 2009. Repertoire size and composition in Great Tits: a flexibility test using playbacks. Animal Behaviour 77: 261–269.

Gill SA, Grabarczyk EE, Baker KM, Naghshineh K & Vonhof MJ. 2017. Decomposing an urban soundscape to reveal patterns and drivers of variation in anthropogenic noise. Science of the Total Environment 599: 1191–1201.

Gill SA, Job JR, Myers K, Naghshineh K & Vonhof MJ 2015. Toward a broader characterization of anthropogenic noise and its effects on wildlife. Behavioral Ecology 26: 328–333.

Goodwin SE & Podos J. 2013. Shift of song frequencies in response to masking tones. Animal Behaviour 85: 435–440.

Goodwin SE. & Shriver WG. 2011. Effects of traffic noise on occupancy patterns of forest birds. Conservation Biology 25: 406–411.

Grabarczyk EE, Vonhof MJ & Gill SA. 2020. Social context and noise affect within and between male song adjustments in a common passerine. Behavioral Ecology 31: 1150–1158.

Grade AM & Sieving KE. 2016. When the birds go unheard: highway noise disrupts information transfer between bird species. Biology Letters 12: 20160113.

Graham BA, Sandoval L, Dabelsteen T & Mennill DJ. 2017. A test of the Acoustic Adaptation Hypothesis in three types of tropical forest: degradation of male and female Rufous-and-white Wren songs. Bioacoustics 26: 37–61.

Haftorn S. 1971. Norges fugler. Universitetsforlaget, Oslo, Norway.

Halfwerk W, Bot S, Buikx J, van der Velde M, Komdeur J, ten Cate C & Slabbekoorn H. 2011a. Low-frequency songs lose their potency in noisy urban conditions. PNAS 108: 14549–14554.

Halfwerk W, Bot S. & Slabbekoorn H. 2012. Male Great Tit song perch selection in response to noise-dependent female feedback. Functional Ecology 26: 1339–1347.

Halfwerk W, Holleman LJM, Lessells CM & Slabbekoorn H. 2011b. Negative impact of traffic noise on avian reproductive success. Journal of Applied Ecology 48: 210–219.

Halfwerk W & Slabbekoorn H. 2009. A behavioural mechanism explaining noise-dependent frequency use in urban birdsong. Animal Behaviour 78: 1301–1307.

Hamao S, Watanabe M & Mori Y. 2011. Urban noise and male density affect songs in the Great Tit Parus major. Ethology Ecology & Evolution 23: 111–119.

Handford P & Lougheed SC. 1991. Variation in duration and frequency characters in the song of the Rufous-collared Sparrow, Zonotrichia capensis, with respect to habitat, trill dialects and body size. Condor 93: 644-658.

Handley HG & Nelson DA. 2005. Ecological and phylogenetic effects on song sharing in songbirds. Ethology 111: 221-238.

Hansen BT & Slagsvold T. 2004. Early learning affects social dominance: interspecifically cross-fostered tits become subdominant. Behavioral Ecology 15: 262-268.

Hartshorne C 1973. Born to sing. An interpretation and world survey of bird song. Indiana University Press, Bloomington.

Huffeldt NP & Dabelsteen T. 2013. Impact of a noise-polluted urban environment on the song frequencies of a cosmopolitan songbird, the Great Tit (Parus major), in Denmark. Ornis Fennica 90: 94–102.

Hunter ML. 1980. Microhabitat selection for singing and other behaviour in Great Tits, Parus major: Some visual and acoustical considerations. Animal Behaviour 28: 468–475.

Hunter ML & Krebs JR. 1979. Geographic variation in the song of the Great Tit (Parus major) in relation to ecological factors. Journal of Animal Ecology 48: 759–785.

Husby M. 2017. Colour aberrations in Eurasian Magpies Pica pica in Europe. Ornithological Science 16: 111–119.

IBM. 2021. IBM SPSS Statistics Armonk, NY: IBM Corp.

Irwin DE. 2000. Song variation in an avian ring species. Evolution 54: 998–1010.

Irwin DE, Bensch S & Price TD. 2001. Speciation in a ring. Nature 409: 333-337.

Kaluthota C, Brinkman BE, dos Santos EB & Rendall D. 2016. Transcontinental latitudinal variation in song performance and complexity in House Wrens (Troglodytes aedon). Proc R Soc B 283: 20152765.

Katti M & Warren PS. 2004. Tits, noise and urban bioacoustics. Trends in Ecology & Evolution 19: 109–110.

Krebs JR. 1977. Song and territory in the Great tit Parus major. Pp. 47-62 in: Stonehouse B & Perrins C (Eds.) Evolutionary Ecology. Palgrave, London.

Krebs JR, Ashcroft R & Webber M. 1978. Song repertoires and territory defence in the Great Tit. Nature 271: 539–542.

Krebs JR, Avery M & Cowie RJ. 1981. Effect of removal of mate on the singing behavior of Great Tits. Animal Behaviour 29: 635–637.

Kroodsma DE. 1982. Song repertiores: problems in their definition and use. Pp. 125–146 in Kroodsma DE, Miller EH & Oullet H (Eds.) Acoustic communication in birds. Volume 2. Song learning and its consequences. Academic Press, New York.

Kunc HP & Schmidt R. 2021. Species sensitivities to a global pollutant: A meta-analysis on acoustic signals in response to anthropogenic noise. Global Change Biology 27: 675–688.

Lampe HM, Larsen ON, Pedersen SB & Dabelsteen T. 2007. Song degradation in the hole-nesting Pied Flycatcher Ficedula hypoleuca: Implications for polyterritorial behaviour in contrasting habitat-types. Behaviour 144: 1161–1178.

Lehtonen L. 1983. The changing song patterns of the Great Tit Parus major. Ornis Fennica 60: 16–21.

Lo S & Andrews S. 2015. To transform or not to transform: using generalized linear mixed models to analyse reaction time data. Frontiers in Psychology 6: 1–16.

Lombard E. 1911. Le signe de l’élévation de la voix. Ann. Malad. l’Oreille Larynx 37: 101–119.

Mace R. 1987. Why do birds sing at dawn? Ardea 75: 123–132.

Marler P & Tamura M. 1964. Culturally transmitted patterns of vocal behavior in sparrows. Science 146: 1483–1486.

McGregor PK, Dabelsteen T, Shepherd M & Pedersen SB. 1992. The signal value of matched singing in great tits: evidence from interactive playback experiments. Animal Behaviour 43: 987–998.

McGregor PK & Krebs JR. 1982. Song types in a population of Great Tits (Parus major): their distribution, abundance and acquisition by individuals. Behaviour 79: 126–152.

McGregor PK & Krebs JR. 1989. Song learning in adult Great Tits (Parus major): effects of neighbours. Behaviour 108: 139–159.

McGregor PK, Krebs JR & Perrins CM. 1981. Song repertoires and lifetime reproductive success in the Great Tit (Parus major). American Naturalist 118: 149–159.

Mendes S, Colino-Rabanal VJ & Peris SJ. 2017. Acoustic adaptation in Turdus leucomelas (Passeriformes: Turdidae) songs to different levels of anthropogenic noise, in the metropolitan area of Belem, Para, Brazil. Revista De Biologia Tropical 65: 633–642.

Mockford EJ & Marshall RC. 2009. Effects of urban noise on song and response behaviour in great tits. Proc R Soc B 276: 2979–2985.

Mockford EJ, Marshall RC & Dabelsteen T. 2011. Degradation of rural and urban Great Tit song: testing transmission efficiency. Plos One 6: e28242.

Moks K, Tilgar V, Thomson RL, Calhim S, Järvisto E, Schuett W, Velmala W & Laaksonen T. 2016. Predator encounters have spatially extensive impacts on parental behaviour in a breeding bird community. Proc R Soc B 283: 20160020.

Morton ES. 1975. Ecological sources of selection on avian sounds. American Naturalist 109: 17–34.

Mundry R. 2011. Issues in information theory-based statistical inference-a commentary from a frequentist’s perspective. Behavioral Ecology and Sociobiology 65: 57–68.

Najar N & Benedict L. 2019. The relationship between latitude, migration and the evolution of bird song complexity. Ibis 161: 1–12.

Pohl NU, Leadbeater E, Slabbekoorn H, Klump GM & Langemann U. 2012. Great Tits in urban noise benefit from high frequencies in song detection and discrimination. Animal Behaviour 83: 711–721.

Pohl NU, Slabbekoorn H, Klump GM & Langemann U. 2009. Effects of signal features and environmental noise on signal detection in the Great Tit, Parus major. Animal Behaviour 78: 1293–1300.

Reijnen R & Foppen R. 1991. Effect of road traffic on the breeding site tenacity of male Willow Warblers (Phylloscopus trochilus). Journal für Ornithologie 132: 291–295.

Reijnen R & Foppen R. 1994. The effects of car traffic on breeding bird populations in woodland. 1. Evidence of reduced habitat quality for Willow Warblers (Phylloscopus trochilus) breeding close to a highway. Journal of Applied Ecology 31: 85–94.

Reijnen R & Foppen R. 1995. The effects of car traffic on breeding bird populations in woodland .4. Influence of population-size on the reduction of density close to a highway. Journal of Applied Ecology 32: 481–491.

Reijnen R, Foppen R & Meeuwsen H. 1996. The effects of traffic on the density of breeding birds in Dutch agricultural grasslands. Biological Conservation 75: 255–260.

Reijnen R, Foppen R, Terbraak C & Thissen J. 1995. The effects of car traffic on breeding bird populations in woodland. 3. Reduction of density in relation to the proximity of main roads. Journal of Applied Ecology 32: 187–202.

Rios-Chelen AA, Cuatianquiz-Lima C, Bautista A & Martinez-Gomez M. 2018. No reliable evidence for immediate noise-induced song flexibility in a suboscine. Urban Ecosystems 21: 15–25.

Rios-Chelen AA, Lee GC & Patricelli GL. 2015. Anthropogenic noise is associated with changes in acoustic but not visual signals in red-winged blackbirds. Behavioral Ecology and Sociobiology 69: 1139–1151.

Rios-Chelen AA, Quiros-Guerrero E, Gil D & Garcia CM. 2013. Dealing with urban noise: Vermilion Flycatchers sing longer songs in noisier territories. Behavioral Ecology and Sociobiology 67: 145–152.

Ripmeester EAP, 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. Behavioral Ecology and Sociobiology 64: 409–418.

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.

Rivera-Gutierrez HF, Matthysen E, Adriaensen F & Slabbekoorn H. 2010. Repertoire sharing and song similarity between Great Tit males decline with distance between forest fragments. Ethology 116: 951–960.

Rivera-Gutierrez HF, Pinxten R & Eens M. 2011. Difficulties when assessing birdsong learning programmes under field conditions: a re-evaluation of song repertoire flexibility in the Great Tit. Plos One 6: e16003.

Rønning OI. 1972. Vegetasjonslære. Universitetsforlaget, Oslo.

Salaberria C & Gil D. 2010. Increase in song frequency in response to urban noise in the Great Tit Parus major as shown by data from the Madrid (Spain) city noise map. Ardeola 57: 3-11.

Sasvari L & Orell M. 1992. Breeding success in a north and a Central-European population of the Great Tit Parus major. Ornis Scandinavica 23: 96–100.

Singh P & Price TD. 2015. Causes of the latitudinal gradient in birdsong complexity assessed from geographical variation within two Himalayan warbler species. Ibis 157: 511–527.

Slabbekoorn H. 2013. Songs of the city: noise-dependent spectral plasticity in the acoustic phenotype of urban birds. Animal Behaviour 85: 1089–1099.

Slabbekoorn H & den Boer-Visser A. 2006. Cities change the songs of birds. Current Biology 16: 2326–2331.

Slabbekoorn H, Ellers J & Smith TB. 2002. Birdsong and sound transmission: the benefits of reverberations. Condor 104: 564–573.

Slabbekoorn H & Peet M. 2003. Birds sing at a higher pitch in urban noise – Great Tits hit the high notes to ensure that their mating calls are heard above the city’s din. Nature 424: 267–267.

Slagsvold T, Dale S & Saetre GP. 1994. Dawn singing in the Great Tit (Parus major) - mate attraction, mate guarding, or territorial defense. Behaviour 131: 115–138.

Snijders L, van Oers K & Naguib M. 2017. Sex-specific responses to territorial intrusions in a communication network: Evidence from radio-tagged Great Tits. Ecology and Evolution 7: 918–927.

SPSS. 2005. Linear Mixed-Effects Modeling in SPSS: An introduction to the MIXED procedure. In SPSS Techical report ( pp. 1–29.

Stone E. 2000. Separating the noise from the noise: A finding in support of the „Niche Hypothesis,” that birds are influenced by human-induced noise in natural habitats. Anthrozoos 13: 225–231.

Tietze DT. 2018. Bird species. How they arise, modify and vanish. Springer, Cham, Switzerland.

Tietze DT, Martens J, Fischer BS, Sun YH, Klussmann-Kolb A & Packert M. 2015. Evolution of leaf warbler songs (Aves: Phylloscopidae). Ecology and Evolution 5: 781–798.

Tolvanen J, Seppanen JT, Monkkonen M, Thomson RL, Ylonen H & Forsman JT. 2018. Interspecific information on predation risk affects nest site choice in a passerine bird. BMC Evolutionary Biology 18: art181.

Warren PS, Katti M, Ermann M & Brazel A. 2006. Urban bioacoustics: it’s not just noise. Animal Behaviour 71: 491–502.

Weir JT & Wheatcroft D. 2011. A latitudinal gradient in rates of evolution of avian syllable diversity and song length. Proc R Soc B 278: 1713–1720.

Wilkins MR, Seddon N & Safran RJ. 2013. Evolutionary divergence in acoustic signals: causes and consequences. Trends in Ecology & Evolution 28: 156–166.

Zbyryt A, Mikula P, Ciach M, Morelli F & Tryjanowski P. 2021. A large-scale survey of bird plumage colour aberrations reveals a collection bias in Internet-mined photographs. Ibis 163: 566–578.

Zuur AF, Ieno EN & Elphick CS. 2010. A protocol for data exploration to avoid common statistical problems. Methods in Ecology and Evolution 1: 3–14.

Great Tit male. Photo: Alf Tore Mjøs.




How to Cite

Husby, M., & Slagsvold, T. (2023). The number of syllables per phrase in Great Tit (Parus major) song decreases in strong anthropogenic noise and at northern latitudes. Ornis Norvegica, 46, 28–42.