Paper, Roll with the fear: environment and state dependence of Pill Bugs

Most studies on animal personality evaluate individual mean behaviour to describe individual behavioural strategy, while often neglecting behavioural variability on the within-individual level. However, within-individual behavioural plasticity (variation induced by environment) and within-individual residual variation (regulatory behavioural precision) are recognized as biologically valid components of individual behaviour, but the evolutionary ecology of these components is still less understood. Here, we tested whether behaviour of common pill bugs (Armadillidium vulgare) differs on the among- and within-individual level and whether it is affected by various individual specific state-related traits (sex, size and Wolbachia infection). To this aim, we assayed risk-taking in familiar vs. unfamiliar environments 30 times along 38 days and applied double modelling statistical technique to handle the complex hierarchical structure for both individual-specific trait means and variances. We found that there are significant among-individual differences not only in mean risk-taking behaviour but also in environment- and time-induced behavioural plasticity and residual variation. Wolbachia-infected individuals took less risk than healthy conspecifics; in addition, individuals became more risk-averse with time. Residual variation decreased with time, and individuals expressed higher residual variation in the unfamiliar environment. Further, sensitization was stronger in females and in larger individuals in general. Our results suggest that among-individual variation, behavioural plasticity and residual variation are all (i) biologically relevant components of an individual’s behavioural strategy and (ii) responsive to changes in environment or labile state variables. We propose pill bugs as promising models for personality research due to the relative ease of getting repeated behavioural measurements.

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