The correlation between colouration and exploration behaviour varies across hierarchical levels in a wild passerine bird

IIn vertebrates, darker individuals are often found to be more active and willing to take risks (representing characteristics of a ‘proactive’ coping style), whereas lighter individuals are instead more cautious and less active (representing characteristics of a ‘reactive’ coping style). It is thus generally expected that melanin-based colouration and proactivity form a suite of positively integrated traits at the among-individual level. Here we use a multi-generational pedigree of free-living great tits (Parus major) to partition variation in, and the correlation between, melanin-based breast stripe (‘tie’) size and exploration behaviour (a proxy for coping style) into its among- and within-individual components. We show that both traits harbour heritable variation. Against predictions, tie size and speed of exploration were negatively correlated at the among-individual level due to the combined influences of permanent environmental and additive genetic effects. By contrast, the two tr...