Causes of variation of darkness in flocks of starlings, a computational model

A. Costanzo*, H. Hildenbrandt, C. K. Hemelrijk

*Corresponding author for this work

Research output: Contribution to journalArticleAcademicpeer-review

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The coordinated motion of large flocks of starlings is fascinating for both laymen and scientists. During their aerial displays, the darkness of flocks often changes, for instance dark bands propagate through the flock (so-called agitation waves) and small or large parts of the flock darken. The causes of dark bands in agitation waves have recently been shown to depend on changes in orientation of birds relative to the observer rather than changes in density of the flock, but what causes other changes in darkness need to be studied still and this is the aim of the present investigation. Because we cannot empirically relate changes in darkness in flocks to quantities, such as position and orientation of the flock and of its members relative to the observer, we study this in a computational model. We use StarDisplay, a model of collective motion of starlings, because its flocks resemble empirical data in many properties, such as their three-dimensional shape, their manner of turning, the correlation of heading of its group-members, and its internal structure regarding density and stability of neighbors. We show that the change in darkness in the flocks perceived by an observer on the ground mostly depends on the observer’s distance to the flock and on the degree of exposure of the wing surface of flock members to the observer, and that darkness appears to decrease when birds roll during sharp turns. Remarkably, the darkness of the flock perceived by the observer was neither affected by the orientation of the flock relative to the observer nor by the density of the flock. Further studies are needed to investigate changes in darkness for flocks under predation.

Original languageEnglish
Pages (from-to)91-105
Number of pages15
JournalSwarm intelligence
Early online date25-Nov-2021
Publication statusPublished - Jun-2022


  • Collective motion
  • Computational modeling
  • Darkness in flocks
  • Flocking of birds
  • Video analysis

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