Journal of Ornithology

, Volume 159, Issue 2, pp 389–400 | Cite as

Song structure and variability in the Black-headed Bunting (Emberiza melanocephala)

  • Sanja Barišić
  • Davor Ćiković
  • Vesna Tutiš
  • Jelena Kralj
  • Herbert Hoi
Original Article


Fighting and flirting are the main evolutionary forces behind the development of bird song and both contribute to different song characteristics. By comparison of vocalisations throughout bird taxa, we can help to reconstruct the evolutionary history of the song. In this study, we provide the first detailed analysis of song structure and song type variability in the Black-headed Bunting (Emberiza melanocephala). We quantify the fine structural characteristics of the song and describe the song and syllable repertoire size. Further, we describe intraindividual song variability over time and interindividual song variability over space, and we explore whether this species uses syntactic rules to create songs. Our analyses reveal that the Black-headed Bunting has a short yet rather complex song and that the syllable pool is extensive. Its song is constructed of three parts that differ in both complexity and rhythm, suggesting the use of syntactic rules in constructing a song. The male song is typically built of 9–13 syllables, with 7–12 that are non-repetitive. Each male sings only one stereotyped song, which remains identical throughout the male’s lifespan, providing evidence for age-limited song learning. Songs are individually distinct, suggesting the importance of song in individual recognition of males. Two spatial levels of song variation in the Black-headed Bunting are shown. On a microgeographical level, relatively small song neighbourhoods are formed with a handful of birds singing the same song type. On a macrogeographical level, dialects can be distinguished by the end part of a song. The most striking feature of the male Black-headed Bunting song is the immense variability of song types. Typically, only a small number of males shared the same song type, and males with unique song types occurred frequently as well. The high song complexity and single-song repertoire found in the Black-headed Bunting could reflect a high level of polygyny and a low level of parental care, therefore pointing to strong intersexual selection on the male song.


Song dialect Song similarity Single-song species Age-limited song learners 


Aufbau und Variabilität des Gesangs männlicher Kappenammern (Emberiza melanocephala) Kämpfen und Flirten sind zwei wichtige Funktionen des Vogelgesangs und damit auch treibende Kräfte der Evolution, die zur Ausprägung von Vogelgesängen beitragen, dabei aber unterschiedliche strukturelle Merkmale fördern. Um die Evolution von Vogelgesängen zu rekonstruieren, ist unter anderem der Vergleich von Gesängen verschiedener Vogelarten eine wertvolle Methode. In dieser Studie präsentieren wir dazu die erste detaillierte quantitative Analyse und Beschreibung von Struktur und Variabilität des Gesangs männlicher Kappenammer (Emberiza melanocephala). Neben Gesangsaufbau, der Ausprägung von Gesangstypen und der Silbenrepertoiregröße werden intra- und interindividuelle Gesangsunterschiede sowie mögliche zugrundeliegende syntaktische Regeln untersucht. Unsere Analysen zeigen, dass männliche Kappenammern kurze aber komplexe Gesänge produzieren, die sie aus einem umfangreichen Silbenpool schöpfen. Der typische Gesang von Kappenammermännchen besteht aus drei Teilen, die sich in Komplexität und Rhythmus unterscheiden, was auf die Verwendung syntaktischer Regeln beim Gesangsaufbau hindeutet. Eine Gesangstrophe besteht typischerweise aus 9-13 Silben wovon 7-12 nicht wiederholt werden. Jedes Männchen produziert nur einen Strophentyp der über die gesamte Lebensdauer unverändert vorgetragen wird. Die Gesänge unterscheiden sich aber zwischen Männchen und dürften der individuellen Erkennung dienen. Bezüglich geografischer Unterschiede im Gesang zeigen sich gleiche oder ähnlichen Gesangstypen zwischen Nachbarn. Großräumig können Dialekte differenziert werden die sich durch den Endteil einer Strophe unterscheiden. Typisch für den Gesang männlicher Kappenammern ist eine Vielfalt an Strophentypen, wobei meist nur wenige Männchen denselben Strophentyp teilen, viele Männchen aber einzigartige Strophen produzieren. Die Tatsache, dass jedes Männchen nur einen Strophentyp produziert, die einzelnen Strophen aber relativ komplex sind, deuten auf eine hohe Neigung zu Polygynie und eine geringe Bereitschaft der Männchen, sich an der Brutfürsorge zu beteiligen hin. Intersexuelle Selektion dürfte also für den Gesang männlicher Kappenammern von großer Bedeutung sein.



We thank Marko Barišić for making most of the audio recordings, villagers from Dazlina for allowing us to study birds on their land and for making our stay there a very pleasant experience, and especially Zdravko and Anka Muić, who generously welcomed us into their home. The study was partly funded by the Environmental Protection and Energy Efficiency Fund. We are grateful to anonymous reviewers and editors for providing helpful comments that improved the manuscript.

Compliance with ethical standards

Ethics approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed. Bird capture and ringing were performed under a ringing licence (Institute of Ornithology, Croatian Academy of Sciences and Arts) and special authorisation for protected species from the Ministry of Environment and Energy (issued for each year of the study).


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Copyright information

© Dt. Ornithologen-Gesellschaft e.V. 2017

Authors and Affiliations

  1. 1.Institute of OrnithologyCroatian Academy of Sciences and ArtsZagrebCroatia
  2. 2.Department of Integrative Biology and Evolution, Konrad Lorenz Institute of EthologyUniversity of Veterinary Medicine, ViennaViennaAustria

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