Behavioral evidence for two-stage sleep in spiders

Sleep is a fundamental behavioral state found across the animal kingdom, yet most of what we know about sleep is heavily vertebrate-biased. Studies are scarce or nonexistent in all but a few invertebrate lineages. A recent study of an REM sleep-like behavior in a jumping spider has reported unexpected parallels to sleep expression in vertebrates. So far, experimental evidence that these behavioral indicators truly represent sleep was lacking in spiders. Here, we present a comprehensive investigation into sleep behavior in the spider Portia fimbriata (Salticidae). We demonstrate diurnal activity, a free-running circadian rhythm, robust sleep architecture, regularly occurring REM-like behaviors, elevated arousal thresholds during those REM-like states, as well as rebound effects following sleep disruption. We conclude that the behavior we observe in spiders is behavioral sleep, providing evidence for an REM-like sleep substate in a terrestrial invertebrate. These results contribute to fundamental questions about the evolution and function of sleep across animals.