The ecology of sleep in non-avian reptiles

Sleep is ubiquitous in the animal kingdom and yet displays considerable variation in its extent and form in the wild. Ecological factors, such as predation, competition, and microclimate, therefore are likely to play a strong role in shaping characteristics of sleep. Despite the potential for ecological factors to influence various aspects of sleep, the ecological context of sleep in non-avian reptiles remains understudied and without systematic direction. In this review, we examine multiple aspects of reptilian sleep, including (i) habitat selection (sleep sites and their spatio-temporal distribution), (ii) individual-level traits, such as behaviour (sleep postures), morphology (limb morphometrics and body colour), and physiology (sleep architecture), as well as (iii) inter-individual interactions (intra- and inter-specific). Throughout, we discuss the evidence of predation, competition, and thermoregulation in influencing sleep traits and the possible evolutionary consequences of these sleep traits for reptile sociality, morphological specialisation, and habitat partitioning. We also review the ways in which sleep ecology interacts with urbanisation, biological invasions, and climate change. Overall, we not only provide a systematic evaluation of the conceptual and taxonomic biases in the existing literature on reptilian sleep, but also use this opportunity to organise the various ecological hypotheses for sleep characteristics. By highlighting the gaps and providing a prospectus of research directions, our review sets the stage for understanding sleep ecology in the natural world.