Journal of Ornithology

, Volume 158, Issue 1, pp 223–231 | Cite as

A comparison of the diurnal song of the Common Nightingale (Luscinia megarhynchos) between the non-breeding season in The Gambia, West Africa and the breeding season in Europe

  • Silke KipperEmail author
  • Patrick Sellar
  • Clive R. Barlow
Original Article


Understanding the full annual song and life cycle of song birds contributes substantially to our understanding of song mechanisms and functions, as well as behavioural ecology and conservation. However, the singing and behaviour of song birds outside the breeding period have barely been studied; this is particularly true for long-distance migratory species. We analysed recordings of the song of Common Nightingales (Luscinia megarhynchos) obtained from their non-breeding grounds in The Gambia, West Africa and compared these to their song during breeding in Central Europe. With regard to song patterns and singing style, we did not find an obligatory breakdown of song organization during non-breeding, but that the birds may sing a typical and recognizable breeding song, which varied on an individual basis. The duration of songs as well as their versatility did not differ between the two samples. However, we did find a difference in song type delivery, with more songs directly repeated during breeding as compared to non-breeding. These results support a practice function of song, but leave also room for additional interpretations such as a function in territorial interactions or as a ‘market place’ characteristic to adjust song repertoires and song sequencing among males sharing their breeding as well as non-breeding areas.


Song function Non-breeding singing Plastic song Song practice 


Vergleich des Tag-Gesangs der Nachtigall ( Luscinia megarhynchos ) außerhalb der Brutsaison in Gambia, West-Afrika, mit dem Gesang während der europäischen Brutsaison Die vollständige Erforschung des jährlichen Gesangs- und Lebenszyklus von Singvögeln wird substanziell zum Verständnis von Gesangs-Mechanismen und Funktionen sowie zur Verhaltensökologie und zum Schutz dieser Arten beitragen. Bislang sind jedoch, insbesondere für Langstreckenzieher, Gesang und Verhalten außerhalb der Brutsaison nur wenig beforscht worden. Wir analysierten Gesangsaufnahmen von Nachtigallen (Luscinia megarhynchos) während ihres Aufenthaltes in Gambia, West-Afrika, und verglichen diese Gesänge mit den Taggesängen während der Brutzeit in Zentral-Europa. Weder die Strophentypen noch die Struktur des Gesanges zeigten einen obligaten „Zusammenbruch“ außerhalb der Brutsaison. Stattdessen sangen alle analysierten Vögel Voll-Gesang, wie er während der Brutsaison typisch ist, allerdings zu individuell sehr unterschiedlichen Anteilen. Die Dauer der Strophen sowie die Gesangsversatilität unterschieden sich nicht zwischen den beiden Kohorten. Allerdings zeigte sich ein Unterschied in der sequenziellen Organisation: während der Brutsaison wiederholten Vögel Strophen häufiger als außerhalb der Brutsaison. Diese Ergebnisse können als Indizien für eine „Einübe-Funktion“ des Gesangs außerhalb der Brutsaison gedeutet werden, allerdings lassen sie auch Raum für weitere Interpretationen wie z. B. territoriale Funktionen oder eine Rolle des Gesangs zum Angleichen der Strophentypen und Gesangssequenzen zwischen Männchen, die in gleichen Brutgebieten leben.



C. R. B. expresses thanks to the Director of the Department of Parks and Wildlife Management, The Gambia for permission to work in protected areas—we look forward to future studies and co-operation! Mawdo Jallow and Lamin Sanyang of the Department of Parks and Wildlife Management assisted in the field and are always a pleasure to work with. Tony Fulford gave freely of his time on several occasions to trap, age and colour-ring Nightingales; we also thank him for interesting discussions. Oliver Fox contributed beneficially to survey efforts and shared good times investigating nocturnal vocal activities. Lawrence Williams, Linda English and staff at Makasutu provided boundless hospitality on multiple occasions. Ringing in Berlin and Brandenburg was done by K. Mortega, C. Bartsch, S. Kiefer and M. Weiss with the permission of the Senatsverwaltung für Stadtentwicklung und Umweltschutz Berlin or the Landesumweltamt Brandenburg, and on behalf of the Vogelwarte Radolfzell (Beringungszentrale an der Max-Planck-Forschungsstelle für Ornithologie) or the Vogelwarte Hiddensee. We are most grateful to M. Weiss, S. Kiefer, I. Adam and C. Bartsch for assistance and critical input during fieldwork and song analysis, and to H. Hultsch and D. Todt for innumerable valuable discussions on Nightingale song. We are very thankful to Peter Slater and Valentin Amrhein for providing valuable comments on an earlier version of the manuscript. All recordings and procedures complied with the current laws of The Gambia and Germany.

Supplementary material

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Supplementary material 1 (XLSX 10 kb)


  1. Amrhein V, Erne N (2006) Dawn singing reflects past territorial challenges in the winter wren. Anim Behav 71:1075–1080CrossRefGoogle Scholar
  2. Amrhein V, Korner P, Naguib M (2002) Nocturnal and diurnal singing activity in the nightingale: correlations with mating status and breeding cycle. Anim Behav 64:939–944CrossRefGoogle Scholar
  3. Apfelbeck B, Kiefer S, Mortega K, Goymann W, Kipper S (2012) Testosterone affects song modulation during simulated territorial intrusions in male black redstarts (Phoenicurus ochruros). PLoS One 7(12):e52009. doi: 10.1371/journal.pone.0052009 CrossRefPubMedPubMedCentralGoogle Scholar
  4. Ball GF, Auger CJ, Bernard DJ, Charlier TD, Sartor JJ et al (2004) Seasonal plasticity in the song control system—multiple brain sites of steroid hormone action and the importance of variation in song behavior. In: Zeigler HP, Marler P (eds) Behavioral neurobiology of birdsong. New York Academy of Sciences, New York, pp 586–610Google Scholar
  5. Bartsch C, Wenchel R, Kaiser A, Kipper S (2014) Singing onstage: female and male common nightingales eavesdrop on song type matching. Behav Ecol Sociobiol 68:1163–1171. doi: 10.1007/s00265-014-1727-6 CrossRefGoogle Scholar
  6. Bartsch C, Weiss M, Kipper S (2015) Multiple song features are related to paternal effort in common nightingales. BMC Evol Biol 15:115. doi: 10.1186/s12862-015-0390-5 CrossRefPubMedPubMedCentralGoogle Scholar
  7. Brumm H, Hultsch H (2000) Pattern amplitude is related to pattern imitation during the song development of nightingales. Anim Behav 61:747–754CrossRefGoogle Scholar
  8. Catchpole CK, Slater PJB (2008) Bird song: biological themes and variations, 2nd edn. Cambridge University Press, CambridgeGoogle Scholar
  9. Erne N, Amrhein V (2008) Long-term influence of simulated territorial intrusions on dawn and dusk singing in the winter wren: spring versus autumn. J Ornithol 149:479–486CrossRefGoogle Scholar
  10. Geberzahn N (2003) Is quantity of song type use in adult birds related to singing during development? Behaviour 140:593–602CrossRefGoogle Scholar
  11. Geberzahn N, Hultsch H (2003) Long-time storage of song types in birds: evidence from interactive playbacks. Proc R Soc B Biol Sci 270:1085–1090CrossRefGoogle Scholar
  12. Geberzahn N, Hultsch H, Todt D (2002) Latent song type memories are accessible through auditory stimulation in a hand-reared songbird. Anim Behav 64:783–790CrossRefGoogle Scholar
  13. Greenberg R, Marra PP (2005) The renaissance of migratory bird biology. In: Greenberg R, Marra PP (eds) Birds of two worlds: the ecology and evolution of temperate-tropical migration systems. Johns Hopkins University Press, Baltimore, pp 437–444Google Scholar
  14. Hahn S, Amrhein V, Zehtindijev P, Liechti F (2013) Strong migratory connectivity and seasonally shifting isotopic niches in geographically separated populations of a long-distance migrating songbird. Oecologia 173:1217–1225. doi: 10.1007/s00442-013-2726-4 CrossRefPubMedGoogle Scholar
  15. Hahn S, Emmenegger T, Lisovski S, Amrhein V, Zehtindjiev P, Liechti F (2014) Variable detours in long-distance migration across ecological barriers and their relation to habitat availability at ground. Ecol Evol 4:4150–4160. doi: 10.1002/ece3.1279 CrossRefPubMedPubMedCentralGoogle Scholar
  16. Hultsch H (1989) Ontogeny of song patterns and their performance mode in nightingales. In: Erber J, Menzel R, Pflüger HJ, Todt D (eds) Neural mechanisms of behaviour. Thieme, Stuttgart, p 113Google Scholar
  17. Hultsch H, Todt D (2001) Developmental trajectories of complex signal systems in animals. The model of bird song. In: Weissenborn J, Höhle B (eds) Approaches to bootstrapping. Phonological, lexical, syntactical and neurophysiological aspects of early language acquisition, vol 2. Benjamins, Amsterdam, pp 309–331CrossRefGoogle Scholar
  18. Hultsch H, Todt D (2008) Comparative aspects of song learning. In: Zeigler HP, Marler P (eds) Neuroscience of birdsong. Cambridge University Press, Cambridge, pp 204–216Google Scholar
  19. Kiefer S, Sommer C, Scharff C, Kipper S, Mundry R (2009) Tuning towards tomorrow? Common nightingales Luscinia megarhynchos change and increase their song repertoires from the first to the second breeding season. J Avian Biol 40:231–236CrossRefGoogle Scholar
  20. Kiefer S, Sommer C, Scharff C, Kipper S (2010) Singing the popular songs? Nightingales share more song types with their breeding population in their second season than in their first. Ethology 116:619–626Google Scholar
  21. Kipper S, Mundry R, Hultsch H, Todt D (2004) Long-term persistence of song performance rules in nightingales (Luscinia megarhynchos): a longitudinal field study on repertoire size and composition. Behaviour 141:371–390CrossRefGoogle Scholar
  22. Kunc HP, Amhrein V, Naguib M (2007) Vocal interactions in common nightingales (Luscinia megarhynchos): males take it easy after pairing. Behav Ecol Sociobiol 61:557–563CrossRefGoogle Scholar
  23. Landys MM, Goymann W, Schwabl I, Trapschuh M, Slagsvold T (2010) Impact of season and social challenge on testosterone and corticosterone levels in a yearround territorial bird. Horm Behav 58:317–325CrossRefPubMedGoogle Scholar
  24. Reudink MW, Marra PP, Kyser TK, Boag PT, Langin KM, Ratcliffe L (2009) Non-breeding season events influence sexual selection in a long-distance migratory bird. Proc R Soc B 276:1619–1626CrossRefPubMedPubMedCentralGoogle Scholar
  25. Sorensen MC, Jenni-Eiermann S, Spottiswoode CN (2016) Why do migratory birds sing on their tropical wintering grounds? Am Nat. doi: 10.1086/684681 PubMedGoogle Scholar
  26. Todt D, Geberzahn N (2003) Age-dependent effects of song exposure: song crystallization sets a boundary between fast and delayed vocal imitation. Anim Behav 65:971–979CrossRefGoogle Scholar
  27. Tramontin AD, Brenowitz EA (2000) Seasonal plasticity in the adult brain. Trends Neurosci 23:251–258CrossRefPubMedGoogle Scholar
  28. Vokurkova J, Petruskova T, Reifova R, Kozman A, Morkovsky L, Kipper S, Weiss M, Reif J, Dolata PT, Petrusek A (2013) The causes and evolutionary consequences of mixed singing in two hybridizing songbird species (Luscinia spp.). PLoS One 8(4):e60172. doi: 10.1371/journal.pone.0060172 CrossRefPubMedPubMedCentralGoogle Scholar
  29. Weiss M (2012) From syntax to details: organisational principles and information encoding in the singing of nightingales (Luscinia megarhynchos). PhD thesis, Freie Universität BerlinGoogle Scholar
  30. Weiss M, Hultsch H, Adam I, Scharff C, Kipper S (2014) The use of network analysis to study complex animal communication systems: a study on nightingale song. Proc R Soc B. doi: 10.1098/rspb.2014.0460 PubMedPubMedCentralGoogle Scholar

Copyright information

© Dt. Ornithologen-Gesellschaft e.V. 2016

Authors and Affiliations

  1. 1.Chair of Zoology, School of Life Sciences WeihenstephanTechnische Universität MünchenFreisingGermany
  2. 2.PurleyUK
  3. 3.Birds of the GambiaBrufutThe Gambia

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