Sources of intraspecific variation in sleep behaviour of wild great tits

Ecologists have recently begun to recognize sleep as a behaviour that is important in animal ecology. The first steps have been taken to characterize sleep in free-living birds, but it is unclear to what extent these results can be generalized between species. To describe sleep behaviour in the wild, we videorecorded great tits, Parus major, in their roosting boxes during two consecutive winters and individuals in captivity for comparison. Here, we examined endogenous and exogenous correlates of sleep behaviour in free-living great tits and addressed the potential confounding issues of studying avian sleep in captivity. Like that in blue tits, Cyanistes caeruleus, sleep behaviour in great tits was strongly related to season, and was affected by sex, age and the environment. Although literature suggests relationships between sleep and risk-taking behaviours, possibly arising from stable differences in physiological state, sleep behaviour appeared to be plastic in great tits, and was not predicted by between-individual variation in exploratory tendencies. Captive tits initiated sleep later than wild individuals, even under natural photoperiodic conditions, suggesting that captivity alters timing and duration of sleep in great tits. Long-term repeatability in sleep behaviour was low for all variables, except morning latency (high repeatability) and evening box entry time, evening latency and frequency of awakenings (no detectable repeatability). Variation in sleep behaviour may largely represent within-individual differences in daily sleep requirements. Our study describes how different observable components of sleep are intercorrelated by providing evidence for significant within-individual correlations between sleep behaviours, which represent the integration of plasticity between traits. Consistent with low repeatability, low between-individual correlations suggest substantial behavioural plasticity in sleep, rather than a correlational structure leading to clear sleep 'syndromes'. Our study provides quantitative evidence for the factors producing phenotypic plasticity in behavioural sleep in an ecological context.