Behavioural syndromes, syndrome deviation and the within- and between-individual components of phenotypic correlations: when reality does not meet statistics

Evolutionary mechanisms leading to correlations across different behaviours, called behavioural syndromes, are hard to study, mostly because behavioural syndromes are group/population level phenomena. Recently (Herczeg and Garamszegi Behav Ecol Sociobiol 66:161–169, 2012), we introduced the concept of syndrome deviation that allows the study of behavioural syndromes at the individual level by focusing on the individual deviation from the hypothetical perfect group-level behavioural correlation. Subsequently, Dingemanse et al. (Behav Ecol Sociobiol 66:1543–1548, 2012) emphasized that behavioural syndromes refer to the between-individual component of phenotypic correlations, and only this component is relevant for syndrome deviation. They also recommended mixed models to decompose the between- and within-individual correlations. We agree that separating these components is important, but the proposed approach is impractical to apply for functionally different behaviours because (1) the assumption of constant within-individual correlations is unjustified and (2) different behaviours cannot be measured at the same time. Further, our simulations based on mixed models show that the statistical differentiation between the within- and between-individual components is inefficient when using realistic sample sizes. Until the separation of between- and within-individual correlations is resolved, we recommend alternative approaches for empirical behavioural syndrome research that consider the repeatability of the behaviours and the optimal balance between within- and between-individual sample sizes. Syndrome deviation calculated from phenotypic correlations of traits that are proven to be individual specific, or from the between-individual correlations if possible, is a meaningful metric to describe behavioural consistency and to explain its evolutionary significance.