Yellowhammer ( Emberiza citrinella ) males sing using individual isochronous rhythms and maximise rhythmic dissimilarity with neighbours

The temporal structure of an animal’s vocal output can be cognitively controlled, presenting an interesting aspect for vocal learning species that benefit from diversification. Although birdsong is the most thoroughly studied aspect of animal communication, its rhythms remain largely unknown. Here, we revisit the question of vocal individuality in the songs of male yellowhammers ( Emberiza citrinella ) purely from a rhythmic perspective. Yellowhammers use simple songs composed of two phrases: the initial phrase , with one individual usually having a repertoire of on average two such phrase types, and the dialect phrase containing the dialect that is the same for all males at a given locality. Some of the initial phrases are commonly shared between various males, but their combinations and frequency contours are individually unique and temporally stable. Using focal recordings of 38 known individuals, collected over three years in the same geographic location, we calculated a set of nine temporal indices to describe each song and compared individuals in a permuted discriminant function analysis. Subsequently, we calculated the potential for individual coding for each of these parameters. To assess whether rhythmic similarity may depend on the singers’ proximity, we calculated vocal dissimilarity as the Euclidean distances between each two males, and used Kendall’s correlation. We show that yellowhammer males use individual rhythms, maintained over different phrase types and carried mostly in the inter-onset-interval variability and syllable rate, as in syllables per second, within the song. There were particularly strong rhythmic differences between singers among the closest neighbours, which decreased over the first 600 m. There was no pattern for neighbours beyond this distance. This study is the first to demonstrate the existence of strong individuality based purely on rhythm, as well as rhythmic differentiation from neighbours, in a songbird.