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

In 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 coloration and proactivity form a suite of positively integrated traits at the among-individual level. Here, we use a multigenerational 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 traits were weakly positively correlated within individuals (i.e. individuals increasing in tie size after moult tended to become more explorative). The patterns of among-individual covariance were not caused by correlational selection as we found additive and opposite selection pressures acting on the two traits. These findings imply that testing hypotheses regarding the existence of a 'syndrome' at the among-individual level strictly requires variance partitioning to avoid inappropriate interpretations as the negative 'unpartitioned' phenotypic correlation between exploration and tie size resulted from counteracting effects of within- and among-individual correlations. Identifying sources and levels of (co)variation in phenotypic traits is thus critical to our understanding of biological patterns and evolutionary processes.

Synergistic and antagonistic interaction between different branches of the immune system is related to melanin-based coloration in nestling tawny owls.

When exposed to parasites, hosts often mount energetically expensive immune responses, and this may alter resource allocation between competing life history traits including other components of the immune system. Here, we investigated whether a humoral immune challenge towards a vaccine reduces or enhances the cutaneous immune responses towards an injection of lipopolysaccharid (LPS, innate immunity) and phytohaemagglutinin (PHA, T-cell immunity) in nestling tawny owls in interaction with the degree of plumage melanin-based coloration. The humoral immune challenge enhanced the response to LPS similarly in differently coloured nestlings. In contrast, the same humoral immune challenge enhanced immune response to PHA in dark reddish melanic nestlings while reducing it in pale reddish melanic nestlings. Our results highlight that both antagonistic and synergistic interactions can take place among branches of immune system, and that the sign and magnitude of these interactions can vary with immune responses involved and the degree of melanin-based coloration.